Skip to main content

Full text of "The Gardens' bulletin, Singapore"

See other formats 


. RA Balin hetion mtn e Pei 
Pirtekrs te ee See eee 


w. 


~ FANE SINT de GY se, . 

MAO alr, Po Doth ta Br dnd» BN yn om bs a 

WO sAhelval Satna ren dp eera nnn 
* » A . . Oe 

7 Wigton ue vad : : 

rs : : . , 

oat ote yan Fi , 


+ 
od ee “- Se 
whic a ~-s. 


~ Se Thes, Moe a 
P= Pe tetas —_ 


ee Loe _s 


fF tho Pigg cm, 


“- 
. 7 Wem Anas tomes ete heme 
ws - ea ce wee 

Nebr reira aed ae ee SRN Mah atone 

SE ROS Repeater ee eee rhs 

: eee Mee Mating omy eta, ae 

~ oe ae et Merino oy he Me, = i uf 

te Weewie. ee z Se fae 

~ . : x 0 AA Ll Be ate doy To SON Cy “a ~~ 

‘1 Ahn APR jis toa shen MPs < . 

~ om © ang ante Moy ante LRRD pee Boe Banta AMR Syne Ce te a, % 4 4 

wD IGeR Ahalh Ln Bint me edcchom mt” ital On es Og, irae 

FO Dol VMs eet a 

OMA Le Met &. an * 

“~ the 


Sa ana nn tee meonsmenn rege tenet 
. ~ * id mar. ae ae eae eee ee 
< Pate en Mit eee an tin Mae 
w + a a Sap Oe NT 
Bawa, Ra aa ONO A A A Few i ein bette eters eMen. - nay Sew theese aie. 
Rit ian oinw/ty iA eae erty hatin oie | Se EN 
y Sat ates Bide, See em a, ~wieedinae tere ~—% 4. h~ue ke ee ripe -tateeenea 
a LEAS NR Bin), on, ity 9, Pal ok an te nan adn Nn ee Regan eee ‘ ie artnet 
INI IRA Me Hi 8 cdtntagge iy, warinte tata ere RO era rans gag wet ree tes tends 
PAPAIN Aton NAA Msra stn, Pie iranatea Ret 
Aaviate Natives van oie on, ten 
’ = FPP 


£ 
Pont, Be ODI aN. CARL hy eubudtala ae 
* wD tN tM ahineey ate Palin ici pacth ar. aupentia e 
Piel AOA Pd SP Sey, a 7 

- ha Pe 


-/ 
te etree ere Bea 
eee ete eh ne Se 
telon mM? Ve Ae! Ae ee et aie uit tere ee 
bdssdetpeaeiona tae ee t en ait ly i=tntetpteeieapce eon etait ee 
9 pa ot ie Re ath Minnie Hume nonin -r-e-éti eee en ee 8, 
» Gadi eae tee ee nite andrea ee Mamas ee See Ree he er, Fe he eae tae at a why 
Sa iietdive take ee em SES b As Pale. .emeMom Naa, — Pe ee ee jal eo ete de ~~)? ae 
a iarddietn en aa ae we Seeedbeadiiptede inte a, ee Serene ~ a ee ee — 
w pniaped dia. om a Dal altmeds Tey oe a ee Pal Ae Me em ~ 
mare ees eis ee ee foie ty * 
ou iene for Mets, oe, & 
Pe nee 
ne * 
PM olen Mie, - 


nn Sa 
eiNiine bee e e 
PDN lek mw 


Miia ae, 
Rey a cr att 
PET Pt it ah ate ae 


pet ee ey 
an ae en 
ars Prat 


ee = ve —, ‘= 
RA. 0m Wye Re ta om ~~. 


en 


Library 


Arnold Arboretum 


co 
i 

‘ho My) K 

yes 7. 


Harvard University 


oe 


; 
ie 
F 
; 
aa 
(ts 
ped 
ie 
(~*£ 
~~ 
vd 
4s ‘ 
* ) 


i ae \e he af ene ee ee 


’ _ - s© . ms. ~ 7 
- ’ _ ie =. 
~ > - F . 
. > ' - 
~ - oe 
, 
- ; ‘ ~w By - 
7 ad : ~~ x ‘ 
f * = 
_* a, = ‘ 
a 
. 
, 
—— 
— 
¢ . 
-? 
es 
. 
Pe iat * et ts 
~~ 
a 
4 


hk 


“ s— = s wm 1s, > .3 2. $5 a +, 
\ 
: : Ny 


FE \ REX REA 


ad 


- ~ J ~ — bo. oo J L- > t- > L o~, £ . ~ -, —, 
9 -O.-- -e eea 2H > X 
Ses i i ee eo 2F Kes" PF DEL KE” He Z =z NW} 


CONTENTS 


* 


B. MOLESworTH ALLEN: Dr. R. E. Ho_truM: An Appreciation - - : 


H. M. BurKILL: Richard Eric Holttum, Croix de Guerre. A note - - 
A. C, JeErmy: Dr. R. E. Holttum — Eightieth Celebration - - - - 
K. C. CHEANG & A. G. ALPHONSO: Holttum’s contribution to horticulture in ae 
the Malaysia — Singapore region - - - - - 9-12-39 
BARBARA JOE HOSHIZAKI: Staghorn Ferns Today and Tomorrow - - - 13-15 4 
SOEJATMI SOENARKO: A New Species of Nastus from Sumba - - - 17-19 
Davip W. LEE: On Iridescent Plants - - - - - - 21-29 = 
A. G. Piccotr: The Ferns of Gunong Ulu Kali - - - - 31-43 
G. J. de JONCHEERE: Specific Concept in Humata pectinata - or) STG gpa 45-58 
B. L. Burtr: Curcuma zedoaria - - - - - - - 59-62 
K. IwaTSuKI: Studies in the Systematics of Filmy Ferns II. A note on Meringium 4 
and the taxa allied to this - - - - - - 63-74 
J. DRANSFIELD: Calamus caesius and Calamus trachycoleus Compared - - 75-78 a 
K. U. KRAMER: Synaptospory: a hypothesis. A possible function of spore = 
sculpture in pteridophytes - - - - - - - ito 4 


E. SOEPADMO & E. E. ona ee Motpheioey oe Sage oe Dennsiaciia ae 
sensu Holttum~ - 85.95 


W. R. Pui_ipson: An Enumeration of the Malesian Species of Aralia - - 97-100, ‘2 


LIEW FAH SEONG: Scanning Electron Microscopical Studies on the Spores of aa 


Pteridophytes XI. Oleandraceae - - - - = : 101-110 Bh 
i 122 


R. D. HOoGLAND: Saurauiae Gerontogeae I. Notes on Malayan species 
Tetsuo KoyaMa: The Cyperaceae Tribe Cypereae of Ceylon - - 123-164 
S. C. CHIN: The Limestone Hill Flora of Malaya I - - - ‘ = 165-219 
J. A. CRABBE: Holttum’s New Taxa and Name-changes in Ferns - - - 221- 238 
M. G. Price: Philippine Dryopteris : ; F So eee © 


WARREN HERB WAGNER JR. & FLORENCE S. WAGNER: Fertile-sterile Leaf 
Dimorphy in Ferns - - - - : a : . 


GUNNAR SEIDENFADEN: A note on Dendrobium serpens - - - - 


BENJAMIN C. STONE: Notes on the Systematy of Malayan Phanerogams XXV. 


Araliaceae - - - - - - - - - 
A. C. JermMy & T. G. WALKER: A note on the cytology ba Batrychiags 
lanuginosum and the occurence of genus in Malesia - 2 
W. R. PHILIPSON: A revision of the Malesian species of Arthrophyllum - - 


‘a aa 
fy fro > m, es Ns 


yeaa 
" i Atoohe i 


A SPECIAL ISSUE TO MARK 
THE EIGHTIETH BIRTHDAY 
OF R. E. HOLTTUM, 1975 


YY 


y 
y 


powyp uig jwwsy <q ydoisojoyd 
siodesulg ‘uspieH suRjog “WINTIegiox{ 94} 32 


SL61 ‘WOLLIOW OMNW AAVHOIYy 


THE 


GARDENS’ BULLETIN 
SINGAPORE 


Vol. XXX 10th October, 1977 


Dr. R. E. HOLTTUM: An Appreciation 
by 
B. MoLeswortH ALLEN 


Los Barrios, Spain 


Dr. Holttum when writing about the classification of the Leptosporangiate 
ferns, most of which in the past were united in the Polypodiaceae, stated that 
“these ‘modern’ ferns are so extremely varied in form and in the details of all 
their parts, that no one man could pretend to a critical knowledge of them all, 
even as to their assignation to genera.” 


It is strange that he should write this, for his amazing faculty for critical 
taxonomic interpretation has enabled him to understand so much about these 
living ferns, both in the field and herbaria. Thus he may well have come as near 
to having just this critical knowledge about which he writes, as any one man may 
possibly ever have. Future pteridologists will have not only the advantage of our 
present-day work, but computers will be used to store much of the knowledge 
that people of Dr Holttum’s age have had to keep in their heads. So although 
there will possibly be a more complete taxonomic and cytological understanding 
of the world’s ferns in the future, this will surely be a combination of man and 
his aids, and not held by one man alone owing to the limit of our brain power. 


The steps that Holttum stands on today were made firm by Carl Christensen 
who brought fern classification into the 20th century, and by the important 
morphological work of Bower and others. In turn each pteridologist was fortunate 
in having the published works of great thinkers of the past who were responsible 
for these higher steps from which Christensen and Copeland worked. 


No doubt Holttum was fortunate in that fate pointed the way at the beginning 
of his career, which started as a palaeobotanist under the guidance of no less a 
scientist than Dr Seward. This gave him an excellent background for the phylogeny 
of the living ferns when he arrived in Singapore and commenced his taxonomic 
work, which he was to continue for the next 40-50 years. Again, the area was a 
fortunate choice; its rich fern flora with so many distinct and unrelated genera 
must have given him much scope for his enquiring mind. The many relic ferns 
growing in their natural environment, gave living examples of evolutionary evidence 
towards a new understanding of the ancestry of different genera and their relation- 
ship. The hot, wet climate of Singapore and the Malay Peninsula, without such 
natural calamities as typhoons and long droughts, was conducive to the continua- 
tion of these primitive types of ferns, many of which were not greatly changed 
from their ancestors. Matonia is one such example of a family once widespread 
in Mezozoic times and some fossils have been found with almost unchanged frond 
shape. The distribution of Matonia pectinata is now extremely limited and is 


1 


2 Gardens’ Bulletin, Singapore — XXX (1977) 


confined to a relatively few hill and mountain ridges in the Malay Peninsula and 
Borneo, growing where the soil is leached and poor, and where it is less likely to 
suffer competition from the surrounding lush tropical plantlife. 


All this was at his doorstep so to speak, and he made the most of it which 
many of us have not done, and no doubt he was helped and guided by Copeland’s 
work on Philippine ferns. One assumes that during these years his fertile mind 
was gathering and storing information on the classification and evolution of the 
vast array of the ferns before him. He also assimilated much knowledge of the 
flowering plants as well. 


Then in Singapore between 1942-46 he was able to correlate this knowledge, 
for being cut off from the outside world he was able to work in the herbarium 
of the Singapore Botanic Gardens, almost without interruption. This resulted in 
the impressive taxonomic publication on the ferns of the Malay Peninsula and 
Singapore. In the course of this work, significant facts came to light which altered 
much of the previously accepted concept of the fern classification of Christensen 
and others. He also pointed out that there were, at least, several families which 
needed far more study especially in field observation, one of these being the 
Thelypteridaceae. He kept Thelypteris more or less together thus apposing Cope- 
land (Genera Filicum 1947) who contended that this generic name was invalid, 
Lastrea being the legimate one. Whilst not actually disagreeing with this, Holttum 
maintained that many changes would have to be made in this group of ferns, 
and until this was done, Thelypteris should stand. As we know, it is a revision 
of this vast and complex family that Holttum himself has undertaken. and from 
his active brain we are seeing these ferns in a new light, and currently a series 
of new combinations and genera are emerging. He has also proposed the conserva- 
tion of the name Thelypteris. 


So the publication of the Ferns of Malaya in 1954 with its appendix con- 
taining cytological notes of a 100 Malaysian ferns by Professor Manton, began a 
new era towards the knowledge and understanding of these tropical ferns. At last 
there was a critical study of this area with up to date name combinations basinyms 
and sufficient synonyms from which one could work. 


As those of us who had followed Christensen, Bower and Copeland (and for 
me, Dr Holloway) in our formative years, this book as well as his papers on 
classification, meant yet another pteridological step upwards. Living in Malaya at 
the time, this publication began a new era for me also. I arrived with a copy of 
Beddome’s “Ferns of British India, Ceylon and the Malay Peninsula” (1892), 
and a separate of Holttum’s ‘““The Common Ferns of Singapore” which had been 
written for the Singapore Naturalist in 1924. (It was almost impossible to obtain 
copies of van Alderwerelt van Rosenburgh’s handbook on the ferns of the Malayan 
Islands). 


The diversity of the ferns with so many unfamiliar genera, not to mention the 
richness of the species of these tropical rain forests, was enough to daunt anyone 
without constant access to herbaria and a botanical library. Ridley’s “‘Ferns of the 
Malay Peninsula” (1926) was very limited in its list of species, whilst many of the 
names used were difficult to trace, so that it helped only in a small way. I found 
that Holttum’s key in his paper together with the notes on the ferns, gave a good | 
introduction to the genera one encountered every day, but the vast number of other 
ferns one saw in the jungle was over-powering. It was a lengthy business, often 
empirical. to try to determine species, usually ending in a deep depression of doubt 
which hardly encouraged intelligent collecting, unless one was a collector by 
instinct So Ferns of Malaya made an enormous difference to those interested in 
the S. E. Asian ferns. 


My association with Dr. Holttum, who was a friend of my husband’s parents, 
has spanned many years, but it differs from most fern botanists in as much as I 
was never long enough in any botanical department to undertake fern research, so 


An appreciation 3 


reluctantly became a field botanist, observing and collecting. Holttum never failed 
to give me encouragement in this, especially when it became tedious. It was 
stimulating to be in the jungle with him, for his field knowledge, not only of 
ferns, but also of the angiosperms, was immense, as was his enthusiasm. An 
interesting plant would be heralded by “a-ha, a-ha”, as he bent to look at it and 
use his lens. On these periodic field trips I learned about tropical botany; he 
pointed out the strange stamens of Lauraceae, and the amount of caulifery in 
these rain forests. He seemed untroubled by the continual heat and high humidity, 
and the rough jungle floor affected him far less than me, even although he had 
suffered an attack of polio after the war. He was an easy houseguest having 
continual good spirits, and only once do I remember that these deserted him. It 
was when he brought the sad news of the death of our mutual friend, A.H.G. 
Alston in 1958. The passing of this gentle person cast a shadow over our trip that 
time, which was to see the extent of Cyathea polypoda on Mt. Kledang near Ipoh, 
where it had been locally common. 


Holttum encouraged _ selective 
collecting, stressing the importance 
of ecological factors, and although 
not, I think, a patient man especially 
with what he considered stupidity, 
he would give painstaking help to 
the student of any branch of botany. 
He also had the gift of being able 


to impact his knowledge easily, cy. Me WWHfP 
which is reflected in many of his ee RS? 
students during his professorship : 12 oN 
at the University of Malaya in 1s 3 wie 
Singapore, who have since become | Aza eZ 
competant botanists. He wrote in x ie ea 
simple language to encourage the \ 3 yp 
beginner and his “Plant Life in Oa ae: 
Malaya” (1954) is an excellent Wie hem 
example of simplicity and erudition. eS 
The output of work is staggering a 
when one realises that apart from “ff WS 
the Ferns of Malaya he published Grammitis 
a companion volume on the orchids; holttumii 


a horticultural textbook “‘Gardening 

in the Lowlands of Malaya” (1953) as well as the monographs on gingers and 
bamboos and the current one on the Thelypteridaceae. Any one of these could be 
considered the culmination of a life study. 


Mention must be made of the enormous help given to the field botanist by 
publications of the vegetation of specialised localities. The Ferns of Mt. Kinabalu 
(1934) by Christensen and Holttum in which many species were described, is one 
of these. I speak from experience, having climbed this exciting mountain, for the 
abundant fern flora, although having close affinities with the Malay Peninsula, 
has interesting endemics and a temperate element from the southern hemisphere. 
So much of this could be appreciated by having read this publication first. 
Although much more is now known of the ferns there, this paper remains important, 
not only in the enumeration of the species, but also for its field notes. 


_ Dr. Holttum’s energy seems unabated and he still travels widely, not only 
in the pursuit of ferns and to receive well deserved honours, but to visit his 
daughter and grandchildren in Australia, which he and his wife have just done. 
So we wish them well with many happy years ahead. 


= Sect ( ane av ii} abs, Gaur ed is Ag 
of: % he! Sr uar eat Laren 
. “Bae Fhe of SEMIS. . co | ages, is x 
to Vind “Yon Sebel ts HB Be ies) 
nA, Twedbiktiic dist dew an Seve et2 ww 2 ete 
brs Yt te door oP INS “At ae edeR Bite eB els \} 
oft vernset fight’ soe Leena Peni BA Ss ie) 
crt reine ) jo lrvem st sd} bits .shorwe: Yo eVaeresy 
#hibh rod ain Dee Perl’ h Tt? ots vd bstéer rey ir . 
Bed of devadif# nov 1 cut) 2eol tela bar Te took ¢ ly 
rer of) ee ari “Tie: — does ( bel . 
erviel terete bh sono Yloowhak 2g ia 
EA obyeroi tl | von bse edt lie 
iw | 1 . hetero, 2irit Sipe otic mre 
atts > toe Ieee Ope oe oo aie a 


y 


sy oak "hse osn al rte! 

: ¢ cece oat} an Sati 
ae rosorkita’ ture AGRE fig 
. Hisisoaed Heer (eae 

sibiggie Bor Dies: La 

. S peepee avr 

Ste mete 


re 
+> at 


: 


he) 


ore 


“ To Ty “ 
jof to Fie ad ia 
oth 
. = 
rm of 4 Whe Ae | tr ae n vy 


7% oi + 


_ fii Hotes ‘s 
a: . 
b> 
i Fy 
It = ; % " 
7. is ! . l 
$ af ; 
NN; : Mis 
~ hi : treet 
ae 
? df? iL ay 
t., rays! Pits ster Rares 


“ay : 


Zs, ISR ATS. 2h AIG tow res 


TEAD ee merivvrade ati: oad. 
Ye MEA RAIS boring oat! SY 


. sede a A 2bicionw ett. nor sal ih et 

Qu riash i ond 

hing im st cmieotem sd) 26 . (Ret) “systole ys 

pte TA ai pst hr tie oat oct? noah noua a 

| ' rane tit 6 too jserter ieee 

iy Ol- rerty eplort eOry1 ONS oily Wy oI it 9 - 

> ion -ons sips Hoa Ae heee Bb (: ICAI HIIQOM 
aii SOs Siow 2S" Ve pherna ae Rye ; 

off oi .maikuem® Soiiney dae pods tS . 

piszeincd ‘valchl att WRIW epee § coh lgi 

S19 imc as Hii OO tao Fase 

AR nbUeaildug’ aks Loa arene Sd | 

Sopehionind | einanss WE axti, ot 


ROTO TH rad pekinese 


a? 7 


Richard Eric Holttum, Croix de Guerre 
A Note 
by 
H. M. BurKILL 


Royal Botanic Gardens, Kew 


Holttum’s autobiographical note in Flora Malesiana Bulletin 28 (1975): 2477- 
500 contains an omission, in part at least self-effacing, but I am sure its honourable 
nature is of sufficient interest to bring it to the attention of his friends and 
colleagues. He refers very briefly to his service in the Friends’ Ambulance Unit 
during the Great War of 1914-18. He went up to St. John’s College, Cambridge, in 
October 1914 and took Class I Honours in the Natural Sciences Tripos, part 1, in 
June, 1916. A normal university curriculum would lead on immediately to the 
completion of the degree course in the following year. But conditions were not 
normal and instead he broke the university timetable to join the Friends’ Ambulance 
Unit for service in France, his university career being completed only after peace 
had been restored. His service records, preserved in the Library of the Religious 
Society of Friends in London run: date of joining 12/7/1916; training at Jordans 
12/7-1/8/1916; ward orderly, St. George’s Hospital, London, 3/8-16/12/1916; in 
France: posted to Headquarters Instructional School 2/7/1917; cook Section 
Sanitaire Anglaise 19 27/1/1917; steward, SS.A. 19 25/9/1918; left unit 
7/1/1919. This service with the French Army was recognised by the French 
Government by decoration with La Croix de Guerre, the citation for which reads: 


ECKERSLEY, STANLEY. B.; HEALD, ALFRED; HOLTTUM, R. ERIC; 
JONES, JAMES; WILSON, ANTHONY L.; WILMOT, HERBERT R.; 
WRIGHT, HERBERT 


Ordre du Régiment 20 Janvier 1919 
‘Convoyeurs a la Section Sanitaire Anglaise 19, ont fait preuve en toutes 
circonstances sous le feu de l’ennemi de courage et de dévouement aux 
blessés aux cours des opérations auxquelles cette section a pris part. (Flandres, 
1917; Somme — Mars-Avril, 1918; Marne — Mai, Juin et Juillet, 1918; 
Champagne — Octobre, Novembre, 1918.)’ 


[as in H. Wright: Two Years with the French Army: Section Sanitaire Anglaise 19, 
Pelican Press, London, 1919, p. 117. Also in Meaburn Tatham & James E. Miles 
(Editors): The Friends’ Ambulance Unit, 1914-1919, a record, Swarthmore Press, 
London, 1920, p. 215.] 


The bibliographic list in Flora Malesiana Bulletin 28 gives 1921 as the year of 
his earliest publication. In fact, the period of his service in France in the Friends’ 
Ambulance Unit was the scene for his first published article: Social Life of the 
Convoy, pp. 91-95 in Herbert Wright’s book cited above, predating all others by 
two years. It is an interesting note on how he and his colleagues lived in France, 
and makes light of time that must have been to a very large extent one of tedium, 
squalor, fear and fortitude. The article shows an amateur theatrical group called 
‘The Duds’, but unfortunately he can not be recognised amongst them. 


_ _ Iam grateful to Mr. Malcolm Thomas of the Society of Friends for his help 
in finding this information. 


srt “sh 40%) ccusttto 2 
shot A 

vd > 

aad MA a 


oD vingiod lerat i ™ 

\ a 
Re inivzsintt pol at Stee (sviskqergoidottng 
peas es o2 Jeol M8 Sieg ms Ozu £ A 
t fi wand of tesredai pe Fs 6 

| 3} Viishid yoy St Be a 

f ww oH Rt-DIOE Fo TEES 

al 7ol seni doa bre Sf ik 

] woving lemon A 3 af 

WHS Sore ab-sai jo. ne + 

sit sor se besten f 5 | 

. rf eeth) igs asT4 fii Oo 2 a 

” Sorenée-2ibl wider fi 

nobaod sit ee 


Ly bTBwW 


i ri 
¥ ‘eat vies 
ort td Lr per 
7 
t 2 . 
; nity j OL 
ty } "d ry H be 
+" ? 
F ee cout 
has tet bs Ge ee} | 
i P. 4 Puase. #4 
$2 
ingeniga yr, oh 
: a] : Hood. Fe oe HPO 


“nore | ab BY. sk alee Bee 
run @pogteheo wh wwe sir 
ra, 7 - ; fave’ ih | ort 


Clima vO¥i SVC: of? en! 


ant, opesph Aare st Arh Qaae re 
el muds ap c8h& UEP caer aobirok 
+ QUGL-DMOL Mg Sones “hasty — 
fe ct 
+t savig BE stole Hopieelet ore ai Jail 96 aga 
EB soiNa3 et Io ROL way cli ra 
C. SSE th tog taut eid. sob og 

ai areas ney jood aa 

Bott . Devil. Be alle 4 eid baa oat weed bai 
fie: 1 ano fagtko ay r Ca a of ee wee 
hey k- AF Ui B> 17650) Waite. AR-@ 
cil tepinne begingeret oc. J 


@loiwb zi int 7usci ts OPE IS ie 


P od a we 7 


Dr. R. E. Holttum — Eightieth Celebration 
by 


A. C. JERMY 


British Museum (Natural History) 
London 


On 30 July 1975, some 75 friends and colleagues gathered in the Jodrell 
Laboratory, Royal Botanic Gardens, Kew, England, at the kind invitation of the 
Director, Professor J. H. Heslop-Harisson, to celebrate the eightieth birthday of 
Dr. Richard Eric Holttum. 


The occasion was organised by a group representing the Linnean Society of 
London (from which Dr, Holttum was presented with the Linnean Medal in 
1964), the British Pteridological Society (of which he was a Past President) and 
the Royal Botanic Gardens, Kew, at which Holttum spent most of his retired, but 
hard-working, hours. 


After sherry in the Staff Common Room guests assembled in the Lecture 
Theatre to applaud a number of presentations to Eric Holttum made by 
distinguished guests on behalf of well-wishers both in England and abroad. 
Mr. P. M. Brenan, Deputy Director of R. B. G. Kew, representing the Director 
who was unfortunately abroad, presided over the proceedings. He welcomed the 
guests and presented Dr. Holttum, on behalf of the Director and staff of Kew, 
with a picture of Holttumara ‘Cochineal’ painted by Rosemary Lowe from 
material specially sent from Singapore. 


Professor Irene Manton, President of the Linnean Society of London, then 
presented Dr. Holttum with a portrait of himseif drawn by Mrs. Julia Pannett and 
subscribed to by friends and colleagues in the United Kingdom, Singapore, New 
Guinea, New Zealand, Holland, Denmark, Switzerland, United States of America, 
Jamaica, South Africa and India. Professor Manton went on to present him with 
a special issue of the Fern Gazette, the journal of the British Pteridological Society, 
published on that day in Holttum’s honour. It was then the turn of Dr. G. Seiden- 
faden, formerly Danish Ambassador to Thailand and a well-known orchidologist, 
who rose to read a statement of good wishes from the President, Mr. Quek Kiah 
Huat, and Members of the Orchid Society of South-East Asia, and presented 
Holttum with a gold medal struck by the Society to commemorate his eightieth 
birthday. Telegrams and letters of good wishes were then read from Mr. A. G. 
Alphonso, Deputy Commissioner i/c Singapore Botanic Gardens; the Singapore 
Branch of the Malayan Nature Society; Mr. John Ede, President of the Singapore 
Gardening Society; and Dr. R. M. Tryon, President of the American Fern Society. 

At this point, Mrs. Jean Brenan, presented Mrs. Ursula Holttum with a 
bouquet of flowers, whereupon Professor C.G.J.J. van Steenis, formerly Director 
of the Rijksherbarium, Leiden, on behalf of colleagues at Leiden and the Flora 
Malesiana Foundation, presented Holttum with a live young plant of Platycerium 
holttumii Hennipm. and Joncheere. 

The guests had a pleasant walk through the Gardens to the Restaurant where 
they enjoyed a dinner given in Dr. Holttum’s honour. The guest speakers, 
Mr. Henry Schollick, President of the British Pteridological Society and Professor 
C.G.J.J. van Steenis gave short but amusing speeches, befitting the informality 


7 


8 Gardens’ Bulletin, Singapore — XXX (1977) 


of this delightful and memorable occasion, Eric Holttum, dressed in a colourful 
Singapore shirt, replied both to the presentation before dinner and to the saluta- 
tions of the after-dinner speeches. 


Even the English weather celebrated the occasion: a screen temperature of 
30°C was recorded that day at Kew. 


Guests at 
Professor Holttum’s Birthday Celebrations 
30 July 1975 


Dr C. D. Adams 
Mr G. Black 
Dr D. F. Blaxell 


Mr & Mrs J. P. M. Brenan 


Mr F. H. Brightman 
Mr H. J. Bruty 

Mr & Mrs J. A. Crabbe 
Dr & Mrs J. Croxall 
Dr D. F. Cutler 

Mr J. W. Dyce 

Mr & Mrs P. J. Edwards 
Mr L. Forman 

Dr & Mrs G. W. Gillett 
Dr A. D. Greenwood 
Miss Mary Gregory 

Mr & Mrs J. W. Grimes 
Mr E. Hennipman 

Dr & Mrs G. Herklots 
Miss D. Holttum 

Dr C. Hubbard 

Dr M. Jacobs 

Dr Frances Jarrett 

Mr & Mrs A. C. Jermy 
Mr G. J. de Joncheere 
Mrs R. Lowe 

Dr J. D. Lovis 

Mrs MacDougal 


Prof. Irene Manton 

Mr P. McKenzie Black 

Mr W. Marais 

Dr & Mrs R. Melville 

Dr & Mrs Panigrahi 

Mrs J. Pannett 

Mrs A. P. Passey 

Miss T. K, Power 

Dr & Mrs D. A. Reid 

Prof. & Mrs P. W. Richards 
Dr & Mrs N. Robson 

Mr H. Schollick 

Dr G. Seidenfaden 

Mrs P. Simmonds 

Dr Anne Sleep 

Dr & Mrs W. T. Stearn 
Prof. & Mrs C. G. G. J. van Steenis 
Mr P. Taylor 

Mr P. Temple 

Dr & Mrs B. A. Thomas 
Dr J. Vaughan 

Dr & Mrs T. G. Walker 
Mr P. J. Wanstall 

Mr S. L. Williams 

Dr & Mrs T. C. Whitmore 
Mr & Mrs J. S. Womersley 
Mr & Mrs J. Woodhams 


Holttum’s contribution to horticulture in the 
Malaysia-Singapore region 
by 
K. C. CHEANG* and A. G. ALPHONSO** 


Horticulture both in Malaysiat and Singapore has taken on a very significant 
role to-day. With rapid urbanisation the importance of planting trees and garden 
plants whether for beautification or for other aesthetic reasons, has become not 
only a very important undertaking, but a way of life with the peoples of the 
region. Many, both amateurs and professionals, who are involved with gardening 
and other horticultural pursuits in this region, make constant references to two 
famous publications — “‘Gardening in the lowlands of Malaya,” and ‘Flora of 
Malaya, Vol I — Orchids.’’ The author of these publications is none other than 
Professor R.E. Holttum. To the amateur who tends his house garden, to the 
professional gardener who runs a commercial orchid nursery or ornamental 
garden, to the horticulturists of horticultural establishments, Holttum is a house- 
hold name, That this should be so is not surprising as Professor Holttum has 
done more for the promotion of horticulture in this region than anyone else. 


Holttum first came out to Singapore in 1922, at a time when there were a few 
Chinese commercial gardens growing some orchids and a few flowering and 
foliage plants. Most of these gardens were in Orchard Road, Thomson Road and 
the Newton and Bukit Timah districts of Singapore, and a few in Penang. Interest 
in gardening among the people at this time was keen, but there was no proper 
guidance by an authority on the subject. Holttum befriended the Chinese gardeners, 
and spent considerable time discussing with them horticultural matters, and at the 
same time observing the traditional Chinese methods of growing plants, one of 
which was the use of burnt clay as a potting medium. There was here an inter- 
change of knowledge between the botanist and the practical gardener. A few of 
these gardeners who are alive to-day and whose children now carry on the 
management of their nurseries, speak very highly of Holttum, both as a horti- 
culturist and as a kind man. Holttum’s knowledge of horticulture was not obtained 
from books alone. His was a practical approach to the subject as a result of long 
experience in the growing of tropical plants in the region. He did not as a rule, 
resort wholly to scientific methods, or put forward scientific suggestions. In the 
pursuit of gardening, he always made it easy for the ordinary man. 


In 1928 Holttum together with John Laycock and Emile Galistan, two very 
keen orchid growers founded the Malayan Orchid Society. At this time there was 
a great interest shown by the public in the growing of orchids, and the three must 
have realised that such a Society would play an important role in the promotion 
of orchid growing in the country. A few years later the Singapore Gardening 
Society was formed, and Holttum was mainly responsible for its founding. 
Meetings were regularly held for members, and the two Societies under the expert 
guidance and advice of Holttum flourished, and played a great role in the pro- 
motion of orchid growing and gardening in the region. Annual shows were held, 
the first being an orchid show organised by the Malayan Orchid Society (now 
the Orchid Society of S.E. Asia) on the 27th and 28th of March 1931 at the 


* lately Head, Botanic Gardens, Penang. 
** Botanic Gardens, Singapore. 
ft here denotes Malaya, Sarawak and Sabah. 


10 Gardens’ Bulletin, Singapore — XXX (1977) 


YMCA Building in Stamford Road, Singapore. Holttum was mainly responsible 
for putting up the exhibits of the many species of orchids and the newly raised 
hybrids of the Singapore Botanic Gardens. It was Holttum who for the first time 
at this exhibition demonstrated to the Singaporeans and Malaysians alike, the 
Knudson method of asymbiotic flask culture of growing orchids, showing flasks 
of orchid seeds at the germination stage, others with seedlings at different stages 
of growth, and finally established seedlings planted on pieces of wood. This 
technique in orchid propagation, which Holttum started in 1928 was the turning 
point in the growing of orchids in the Malaysia-Singapore region, and in fact the 
whole of S. E. Asia. Seeds could now be successfully germinated and as such, was 
a tremendous asset in the hybridisation and breeding of orchids in the region. 
Holttum himself produced a large number of hybrids. The orchid shows continued 
annually until 1934, when the first flower show including both ornamental and 
orchid plants was held at the New World Stadium from the 6th-8th April, and 
jointly organised by members of both the Malayan Orchid Society and the 
Singapore Gardening Society. These shows were to continue annually to the present 
time. Holttum must have known that horticultural shows are of immense value 
to both professional and amateur growers, for they serve to raise the standard of 
horticulture in the country by giving the growers a chance to compare their 
exhibits and to show what can be achieved. They also help to encourage the non- 
gardener to take an interest in gardening. Holttum retired as Director of the 
Singapore Botanic Gardens in 1949 and was appointed Professor of Botany at the 
newly formed University of Malaya. He held this post until 1954 when he returned 
to England. For over 25 years he had devoted his time energetically to the 
activities of the Malayan Orchid Society and the Singapore Gardening Society. 


Although the flora of the Malaysia-Singapore area is one of tropical rain 
forest, it is surprising to note that there are few indigenous plants of horticultural 
value. Native ornamental plants and flowering trees are few compared to the 
abundance of species in the flora of the area. The flowering trees and ornamental 
plants grown in one’s garden, in parks and open spaces, by roads and streets and 
even in the Botanic Gardens, have mostly been introduced especially from Central 
and South America, and other parts of Tropical Asia. The introduction of new 
plants into Malaysia and Singapore has therefore played an important role in the 
horticultural beautification of the area. Although private individuals, business 
houses and other establishments have to some extent been responsible in the 
introduction of plants into this region, it was Holttum, during his 27 years of office 
as Assistant Director and Director of the Gardens Department, Straits Settlements 
(Singapore, Malacca and Penang), who was mainly responsible for the countless 
numbers of flowering trees and shrubs introduced to the two Botanic Gardens in 
Singapore and Penang from tropical regions of North and South America, Africa, 
Asia and Australia. Many of these plants to-day are extensively grown in parks, 
by roadsides, and private gardens and commercial nurseries throughout Malaysia 
and Singapore. 

Holttum is a widely travelled person, having attended many international 
horticultural conferences. His lectures and papers presented at these conferences 
included topics on various aspects of horticulture in this region, particularly relating 
to orchids and ferns, 


Some thirty odd plants, both species and hybrids, have been named after 
Holttum. Perhaps the trigeneric orchid hybrid Holttumara cochineal (Arachnis x 
Renanthera x Vanda) is the most outstanding as far as horticultural plants are 
concerned. A painting of this hybrid was made by a staff member of the Royal 
Botanic Gardens, Kew, and presented to Holttum on his 80th birthday. Some 
years back the authors on a plant collecting trip to Kaki Bukit in Perlis, Peninsula ~ 


Zz 


Holttum’s contribution 11 


Malaysia, discovered a staghorn fern which was appropriately named Platycerium 
holttumii, for no one in this region has done more towards popularising the 
growing of ferns as a decorative house plant than Holttum. 

In 1940 for the first time, three local officers were recruited by Holttum for 
horticultural training. K. C. Cheang was assigned to the Botanic Gardens in Penang, 
A. G. Alphonso to the Botanic Gardens in Singapore and N. V. Lange to the Parks 
Department in Singapore and subsequently to the Parks Department in Ipoh, 
Perak. Holttum must have realised at this time that the future of horticulture in 
the region rested with qualified local horticulturists. Up to such time horticulturists 
were expatriate officers recruited from the Royal Botanic Gardens, Kew. Apart 
from the training received directly under Holttum, the officers were sent to the 
College of Agriculture in Serdang, Malaysia, and later K. C. Cheang proceeded to 
Massey University, New Zealand, N. V. Lange to the Institute of Parks. Administra- 
tion, England, and A. G. Alphonso to the Royal Botanic Gardens, Kew. On their 
return they were appointed Curators, and were subsequently to head their 
respective departments. It was during the period of training under Holttum that 
the authors had come to appreciate him as a truly outstanding horticulturist. His 
immense knowledge of local plants and his long standing experience in practical 
gardening, won him the admiration of all who came to know and learn under him. 
Above all, he was a very kind person, and was always willing to help his fellow- 
men. We are proud and honoured to have served under Holttum. 


Holttum’s contribution in the field of horticulture has won him many honours 
and he has been the recipient of a number of awards from horticultural societies 
in the U.S.A., Great Britain and Singapore. In recognition of his work on orchids 
in Malaysia and Singapore, Holttum, at the 4th World Orchid Conference held in 
Singapore in 1963, was awarded the Gold Medal of the Malayan Orchid Society. 
In addition, the Orchid Society of S.E. Asia awards annually a Gold Medal known 
as the Eric Holttum Gold Medal to the originator of the best locally produced 
hybrid. On the occasion of his 80th birthday the Orchid Society of S. E. Asia, 
at a ceremony at the Royal Botanic Gardens, Kew, presented him with a Gold 
Medal suitably inscribed, and through the generosity of the Singapore Gardening 
Society, the Botany Department of the University of Singapore presents annually 
a Holttum Silver Medal to an honours student who has performed well in his 
examinations. 


Perhaps Holttum’s biggest contribution to horticulture in the Malaysia- 
Singapore region lies in his published works. He is a prolific writer, and his many 
articles on gardening in the Malayan Agri-Horticultural Association magazine for 
well over 20 years, are not only instructive but of great benefit to both the amateur 
and professional gardener. The Malayan Orchid Review, the official organ of the 
Malayan Orchid Society was first published in 1931. Holttum was the main contri- 
butor. He was the expert, and his many and varied topics in the Orchid Review 
have helped orchid growing in this region. By far, his best works in horticulture 
and by which the Malaysian and Singaporean will best remember him, are his 
“Gardening in the Lowlands of Malaya” and his ‘Flora of Malaya, Vol I — 
Orchids”, mentioned earlier. The former is an excellent book on_ practical 
gardening in this region. It is as popular to-day as when it was first published in 
1953. The latter, the most outstanding publication on orchids of this area, has 
been a constant companion of the orchid growers. Holttum is also the author 
of many other publications on local plants of horticultural interest. His ‘‘Flora of 
Malaya, Vol Il — Ferns” and articles on Bamboos, Gingers, Maranta, Grasses 
and others, published in the Gardens’ Bulletin, although taxonomic or 
botanical in nature, have in many ways helped the horticulturist in his work, 
especially in the identification of plants he deals with. 


12 Gardens’ Bulletin, Singapore — XXX (1977) 


We in this region owe a debt of gratitude to Holttum for his outstanding 
contribution to horticulture. To-day, in both Malaysia and Singapore, there is a 
thriving orchid industry. The value of present exports of cut orchid blooms from 
both these countries has reached nearly ten million local dollars. Orchid nurseries 
have come up in large numbers, and there is no doubt that the orchid industry will 
continue to expand. This situation would not have come about had it not been 
for the pioneer work of Holttum in orchid growing. His introduction of asymbiotic 
flask culture of orchids n 1928, the breeding and hybridisation of orchids which he 
started, and the encouragement he gave to professionals and amateurs in the field 
of horticulture have borne fruit. 


The role he played as adviser to the Parks Department and in the planting 
of roadside trees in the early years, his able administration of the two Botanic 
Gardens of Singapore and Penang, and the valuable knowledge he has imparted 
to the students as Professor of Botany at the University of Malaya, bear testimony 
to his foresight and vast horticultural knowledge. And above all, his many horti- 
cultural publications will be constant reminders to us all of his dedicated service 
in this region. 

This number of the Gardens’ Bulletin is specially brought out to commemorate 
the 80th birthday of Professor Holttum, and in writing this article the authors join 
the many Malaysians and Singaporeans, the orchid growers, the amateur and 
professional gardeners, and the members of the Orchid Society of S. E. Asia and 
the Singapore Gardening Society, in wishing him good health and happiness in 
the years ahead. 


Staghorn Ferns Today and Tomorrow 


by 
BARBARA JOE HOSHIZAKI 


Botanical Gardens-Herbarium 
University of California 
Los Angeles 


The unique appearance of staghorn ferns (Platycerium) has attracted the 
attention of enthusiasts for well over a hundred years. Modern transportation and 
communication have brought all 18 species into cultivation. So great is interest in 
this fascinating fern group that it seems appropriate to note recent horticultural and 
botanical works that have solved many puzzles and to look ahead to problems yet 
to be answered. 


HorRTICULTURAL CONTRIBUTIONS. Horticultural interest in staghorn ferns has 
greatly benefited botanical work. The perseverence and tenacity of Platycerium 
fanciers who seek out new plants, grow them, and develop new varieties is amazing. 
Through their efforts I have been able to study every one of the known Platycerium 
species as live plants. The opportunity to watch plants grow from spore to maturity 
contributed greatly to the botanical knowledge of the genus. 


Platycerium enthusiasts in United States cultivated a new species from 
Malaysia for several years before it was recognized and named by botanists 
(Joncheere and Hennipman 1970). This handsome plant, now called Platycerium 
holttumii, was well known enough by amateurs to have been photographed for two 
popular books prior to being formally named (Rie et all 1957, Franks 1969). A 
second new species, the P. grande of the Philippines, has been recognized in the 
last few years (Joncheere and Hennipman 1970), another exciting event to 
platycerium fanciers! These relatively recent discoveries kindle the imagination of 
collectors who hope that different species may still be discovered. Collectors continue 
their quest for new and unusual platyceriums. Shipments of P. wallichii have 
reappeared in the United States, and these plants do show some frond variation. 
The significance of these variations has yet to be investigated. These plants were 
reported to have been collected along the India-Burma border. Native plants of 
P. stemaria have an interesting range of frond variation which may interest botanist 
as well (Joe 1964). But besides seeking variations among wild plants, horticulturist 
are developing new garden varieties. The multititude of new garden varieties is due 
to the increased number of people raising plants from spores. A hybrid platycerium, 
P. mentelosii (P. stemaria x superbum) has been developed and other hybrid 
combinations will most likely follow (Hoshizaki 1975). Growers tell us that 
platyceriums are quite plastic in their leaf form. Slight changes in the environment 
may cause base fronds to become partly foliaceous, normally entire fronds to 
become forked and so forth. 


Staghorn fern propagation by meristem or tissue culture is being actively 
investigated by commercial nurseries. They claim that plants may grow twice as 
fast through meristem culture as by spores. Meristem plants tend to produce 
mutations more frequently than spore grown ones and this interest horticulturist 
and geneticist. 


__ The Malaysian plant, Platycerium ridleyi, thought extinct, has been found and 
introduced into United States cultivation. It grows very nicely in southern California 


13 


14 Gardens’ Bulletin, Singapore — XXX (1977) 


greenhouses if adequately protected from slugs and snails. Spores from these 
introduced plants have produced mature plants, thus insuring the species’ survival 
and reducing the need to collect these rare plants from native habitats, a happy 
circumstance for conservationists. 


Much more information is needed on the cultural conditions for Platycerium 
wallichii of Southeast Asia, and P, madagascariense and P. quadridichotomum both 
of Malagasy. These species, being difficult to cultivate, create demands for replace- 
ment, a situation which is not encouraging for conservationists. Hopefully more 
work along the lines of Boyer will help in the successful growth and propagation of 
these plants. Boyer closely examined the ecology, physiology, and mineral needs 
of two African species, P. angolense and P. stemaria (Boyer 1964). 


The type of foregoing activities and results generated by horticulturists hold 
promise of supporting and even giving direction to scientific work and increasing 
our understanding of the genus. To help science, horticulturist should be encouraged 
to maintain reliable records as to where native plants were collected, or if new 
plants were developed in cultivation, the names of the parent or parents involved. 
Conditions which may have caused unusual growth patterns should also be recorded. 
Horticulturists on the other hand should protect native plants and prevent their 
extinction from indiscriminate collecting. 


BOTANICAL CONTRIBUTIONS. The name changes among platycerium species 
confuse and frustrate growers and most botanists. If it is any comfort to know, 
botanist specializing in nomenclature have for years been trying to unravel the 
technical complexities in determining the proper name for certain species. A series 
of papers and rebuttals dealing with this problem has been published in the last 
few years by G. J. Joncheere and the late C. V. Morton (Morton 1964, 1970, 
Joncheere 1967, 1974). The main issues first center around whether Platycerium 
vassei as conceived by botanist, not horticulturist should be called P. alcicorne and 
second, whether P. angolense should be called P. elephantosis, A few botanist are 
following this intricate and complicated problem and their views will be forth- 
coming. Morton’s argument, that the correct spelling of P. stemaria is not P. 
stemmaria, seems to be upheld. The acceptance of P. wandae as the legitimate name 
over P. wilhelminae-reginae- seems without complication (Joncheere 1968). How- 
ever, among plants introduced from New Guinea and now growing at Longwood 
Garden, Pennsylvania are two kinds of P. wandae, One produces long fertile fronds 
fitting the original description of P. wilhelminae-reginae and the other produces 
short fronds closely fitting the description of P. wandae. In all likelihood both 
plants are one species. Short and long fertile frond forms have also been noted in 
P. coronarium, However, further study and field observations on long and short 
frond plants might be of interest. 


It is indeed unfortunate that the name Platycerium grande must now be 
applied to the Philippine plant instead of the Australian plant. The Australian 
plant must now be called P. superbum in accord with Joncheere and Hennipman 
(1970). I examined the rhizome scales of the Philippine plant and they differ from 
the Australian plant, though both are closely related. Rhizome scales of the 
Philippine plant are very similar to P. holttumii except for having slightly fewer 
cells in the marginal hairs (highest number of cells in the longest hairs were mostly 
7 in P. grande and 8 in P. holttumii). The scale morphology is consistent with other 
morphological data which relates P. grande, P. superbum, and P. holttumii 
(Hoshizaki 1970, 1972). 


The relationships of the Australian-Javan species (Platycerium bifurcatum, 
P. veitchii, P. willinckii, and P. hillii) and their distribution still needs to be clarified 
with more data. Some of the features separating these supposedly different species 
are not stable from observations on plants in cultivation. Mr. Ernest Todd of New 
South Wales, Australia has been investigating the distribution of the Australian- 
Javan species. In a personal communication Mr, Todd reports that the collections of 


Staghorn Ferns 15 


P. bifurcatum by Bambler and others in New Guinea were probably taken from 
cultivated or escaped plants most likely introduced by German missionaries in the 
early part of the century. Distributions of some of the Malayan-Asiatic species 
also need further clarification. 


Platycerium coronarium and P. ridleyi are very closely related species, yet 
there is a gap between these two species and their closest relatives. It doesn’t seem 
too far-fetched to speculate that some yet undiscovered species may be found in 
the Malaysian area that bridges this gap and others as well. 


With the relatively wide-spread use of the scanning microscope studies should 
be encouraged on the ontogeny of stellate hairs in this genus and its proposed 
relatives (Pyrrosia and Drymoglossum). It might be worthwhile to also study and 
compare these genera as to sporangial structure. Within the platyceriums are species 
with complete to incomplete annular rings, laterally and apically located lip cells, 
and stalk structure variations. 


Certainly there are many topics I have not touched upon that should be 
considered in future studies. However, I want to stress that botanist and horti- 
culturist have much to look forward to in knowing more about these unusual 
plants. 


Literature cited 


Boyer, Yvette née Menoux. 1964. Contribution a l’estude de l’ecophysiologie de 
deux fougeres epiphytes: Platycerium stemaria (Beauv.) Desv. et P. angolense 
Welw. Théses présentées a la Faculté des Sciences de l'Universite de Paris. 
Série A, No. 4191, No. d’ordre: 5042, 


Franks, Wendy. 1969. Platycerium Fern Facts. Privately published. 119 p. 


Hoshizaki, Barbara Joe. 1970. The Rhizome Scales of Platycerium. Am. Fern J. 
60: 144-160. 


Hoshizaki, Barbara Joe. 1972. Morphology and Phylogeny of Platycerium Species. 
Biotropica 4: 93-117. 


Hoshizaki, Barbara Joe. 1975. A Staghorn Fern (Platycerium) Hybrid. Am. 
Fern J. 65: 99-101. 


Joe, Barbara. 1964. A Review of the Species of Platycerium (Polypodiaceae). 
Baileya 12: 69-124. 


Joncheere, G. J. de. 1967. Notes on Platycerium Desv. I. Nomenclature and 


typification of the genus and species in Desvaux’s original publication of 1827. 
Blumea 15: 441-451. 


Joncheere, G. J. de. 1968. Notes on Platycerium Desv. II. P. wilhelminae-reginae 
v.A.v.R. reduced to P. wandae Rac. Blumea 16: 109-114. 


oe G. J. de. 1974, Nomenclatural notes on Platycerium (Filices) Blumea 
2: 53-55. 


Joncheere, G. J. de and E, Hennipman. 1970. Two new species of Platycerium and 
the identification of P. grande (Fee) Presl. Br. Fern Gazette 10: 113-118. 


en. es V. 1964. The Nomenclature of a Madagascarian Platycerium. Baileya 
12: 36-38. 


perion. C. V. 1970. A further note on the type of Platycerium alcicorne. Am. Fern 
. 60: 7-12. 


Riie, Edgar Aubert de la, Francois Bourliére, Jean Paul Harroy, 1957. The Tropics. 
Knopf. 208p. 


7 


Lhe we ag mainsaiealt ites neaeeity ‘ ee 


\ send 
wath’ 


Aga sett: user Raia 
et 


Palais 
ae See te 


ure nsnanining - 


“oserye rel ta a> 

lay 2 saizege 2 boven laooko cov, o18 Yalu sf 
mosa 1 azopb J, zovitalen toeOl2 ter. bas. Beat 
ni. bauot sd. vem, 29199g2 Do1 piace Wy is, 3502 oe 
Tow es zedio.bng cag, a, fy PER a mf 

blirade ‘eoubule SOSA AIF PT sit jensen elie 7 
Lseecend! er bre aero ic) nt! efierl: onion to eccayodon at 1th 
brid chute gale ot dticwaln oe rigien J Coated ae 
no | alli eaudoue Leeioeengs ag) Sherr: 


intel 


ail baisoo! vilsviqs- bag: yllersish agar 1 Lasts atoige gon, i 
eet ate 7 


on vac t esiegod ete coe 


wr ae Ge miwebh api bartenae 
; , (1 ¢3) Hapa oe sedi ae 
wast 
j : : i 4 
% 4 RATES 
851 
“+ 


Heir gtoipvrih akortceek 3 sino coe 


ih et 
(ie aa SAT Oret 
eel yas PR (aot sg ronA Tei elie 
. ; Ae. oF ’ 
HE ba 
, Mue'e 
; pais 4 - f 7 i se Pe f Ler, 


var tat eS 7 Cis ae 
8 Fy fa : ,. rt ft Thkh oeinit MG ae Sarat 
SF H Bitlet - 4 Tie } - pev2atl (vi ee mat hs DPB ag 


iti PAT AS 


e-santrlot ee AE vee sissies Peale fo rao) 
at fp OF al ay uct # a 


* bored) 

HSH PONIES a, 220 . 
hiss CurTsaviEhkt To? sina Woo’ a Ne ‘air ; 
WHEEL OE went faa aR 1 [} 9b 


fis. 


7 BD ra 4 7 cf ye a ree iy es or 


A new species of Nastus Nees (Gramineae) from Sumba 
by 
SOEJATMI SOENARKO 


Herbarium Bogoriense 
Indonesia 


Summary 


An illustrated description of Nastus reholttumianus S. Soenarko is presented and this new 
species is compared with N. rudimentifer Holtt. and N. obtosus Holtt. 


When Holttum (1956) transferred the Malesian species of Oreiostachys 
Gamble to Nastus Nees, three species of Nastus were known from New Guinea. In 
1967 he described four more species and Bor (1972) added one to these; thus 
there are now eight species endemic to New Guinea. Outside New Guinea there 
is One species; it occurs in Java. 


In most of the Malesian Nastus the spikelets are more or less cylindrical, thus 
the fourth and the fifth glumes, lemma and palea are rounded at the back (except 
those in N. schlechteri (Pilger) Holtt., in which they are more or less 1-keeled), 
with usually prominent nerves; moreover the palea is bifid. A plant collected by 
Mr. Iboet from the island of Sumba (Indonesia) has characteristically small and 
laterally compressed spikelets, lightly 1-keeled glumes, lemma and palea, with 
prominent middle nerve and several (usually 6) faintly lateral nerves, and 
acuminate palea, These structures, together with the leaf size (up to 8 cm long 
and 14 mm wide) differentiate the Sumba plant from the other Malesian Nastus, 
and it is regarded here as a new species, N. reholttumianus. The near relatives of 
this new Nastus are N. rudimentifer Holtt. and N. obtusus Holtt. which have similar 
panicle with spreading branches and short-tipped lower glumes. However, this new 
species differs in several respects from the two latter; their characters are compared 
in the table below: 


characters reholttumianus rudimentifer obtusus 
leaf-blade of flowering | 4.5-8 cm long (8) 10-12 cm long | 12-15 cm long 
branch 7-14 mm wide 15-27 mm wide 12-15 mm wide 
spikelet + laterally + cylindrical + cylindrical 
compressed 
5-7 mm long 12 mm long 13-20 mm long 
rachilla extension absent present, sometimes | usually absent 
absent 
lemma 6 mm long, 10 mm long, 14-17 mm long, with 
glabrous glabrous appressed hairs on the 
back 
palea 
acuminate bifid bifid 


Nastus reholttumianus S. Soenarko, sp. nov., N. rudimentifer Holtt. affinis 
sed spiculis 5-7 mm longis, latere compressis, lemmate 6 mm longo, lamina 8 
longa differt. Typus: Sumba, Iboet 443 (holotypus, BO). 


17 


18 


Fig. 


Gardens’ Bulletin, Singapore — XXX (1977) 


1. Nastus reholttumianus sp. nov. J, flowering branches; 2, spikelet; 3-7, glumes I — 
V respectively; 8, lemma; 9, palea; 10, ovary; 11, lodicule; 12, spikelet after removing 
the glumes and lemma. 


4 

~ 
= 
¢ 


j 


Nastus 19 


Culm not seen, upper part bearing flowering branches 3.5 mm diam., either 
solid or hollow, with annular woody ring below each node; branches glabrous, 
3-8 in each node. Culm-sheath not seen. Flowering branch 11-22 cm long; leaf- 
blades 4.5-8 cm long, 7-14 mm wide, oblong-lanceolate, glabrous, acuminate, 
rounded at the base; leaf-sheath glabrous; ligule less than 1 mm long; auricle up 
to 1 mm long, fringed (fringe 1-2 mm long). Inflorescence open panicle with 
spreading branches, main axis and branches with minute appressed hairs. Spikelet 
more or less laterally compressed, up to 7 mm long, glabrous; glume I: 1.5 mm 
long, narrowly ovate, 1-keeled on the back, with mucronate apex; glume II: 2.5 mm 
ovate, slightly 1-keeled on the back, mucronate; glume III: 3.5 mm long, slightly 
1-keeled on the back, ovate, faintly 6-nerved with conspicuous median nerve (and 
so are the median nerves of glumes I and II); glume IV: 5.5 mm long, ovate to 
ovate-oblong, boat-shaped, acuminate, 3 mm wide, faintly 7-nerved; glume V: 
more or less similar to glume IV, but slightly longer; lemma and palea more or 
less equal, 6 mm long, ovate-oblong, acuminate, boat-shaped to slightly com- 
pressed; lodicule 1 mm long with fringes; ovary stalked with conical fleshy apex, 
glabrous, the stalk often elongates up to 2 mm long; rachilla extension absent; 
stamens 6, anthers 2.5 mm long. (Fig. 1). 


Sumba. Mangiliwari near Maomaru, 9. V. 1925, Iboet 443 (BO, holo). 


The author dedicates this new species to Prof. R. E. Holttum who has 
contributed so much to the knowledge of S. E. Asian bamboos, especially of 
Malaya and New Guinea, and has encouraged the author in studying Malesian 
bamboos. Thanks are due to Sdr. Damhuri (Herbarium Bogoriense) for preparing 
the figures. 


References 


Bor, N. L. (1972). A new species of Nastus from New Guinea. Ost. Bot. Z. 
120: 87-91. 


Holttum, R. E. (1956). The bamboo-genera Nastus and Chloothamnus. Kew. Bull. 
10: 591-594. 


(1967). The bamboos of New Guinea. Kew Bull. 21: 263-292. 


i ert oe 
Sisnimuss pee slosonsl-gnolde ohiw "ee Tt 
qu slanus jgool rot tsd? ee! luail rao fy ‘ds 
dtiw sivinsq b bane sonapesofin! _ Agnok rai She mt: * 
tafodide ctied Uyoeeewqe signiss ita vadborened bay 2ize | Then 
neh at el simuly yuoidel: sack, mint IR of tic Deaws mos Herstal | 

mm COST sinytg eo B 9. CNN hw we ad an) ao bolesdet | ee ih 


-taol nin ra SS 


4 


-sT4 fetta ail mh ? rs witty we ie +e) bee ey ol sth atawpe att tit a. : 5 


cers) ayret mbibaen 20": pou) Hie beyigted vhetat See: aaa ba 
of sieve casob mim 2.2 : Vi stale” bee if acct: 306 he N21 USRA,, 
A emule bayer! sick shiv saan Sani aaa 
SD un ablan Baa sores outer ehgaile idl aan ih 

_ 5 te ri Doerner LEC 1 @oEAI a , yeas Ma tri Rye cant (sin “8 
ni ; wigan , ‘. iw hodlste co iive:, :oRrereiahie Biol EH is alway 
‘jill - ee eet rg pad . St of Gg iy) 2O Rae my Beetay sf 
athe’ — 2 ce rich ae: 2 aa past 1 2 Oag 


aul 


dona Fe 


ety eae 
Folod OF) Les Woah ROSY. cro ket laa h 
i Ode ceuiilolt tt teases Siew blk zoe siheb: hinders : Pepe 
-.  ocoteved: aaa See A¥-ot ove OFF 
Uh ccrchan> ni vesdtume wit, Lomcutlets ‘wkd! BAe esas 
Bis cas rm yOah ARE Nek & ae eilivteat. 2h, OF si ae ane 2) 


S of a .2dnivd! welt Soak Whe Go ee wane i tery) 
iia a. 


19 


* oe i : « : 
Med 
k te 7 at Cae ee 3 
at 2s a ee 7c) ‘Warr 3 sdincdtad t RAY 
, ‘4 
U 
% | 


«& Pal 
ft A 
Pr > 7 
ahs ne 
:7 * 
ros 
“| ke 
Tye,” °? 
a. yy . 
od < 
ee yt J aA 
a he 
7 ie A %, 4 


Moe 
4 
eg * 
ae 


a ae 


sot 


fuming? a. be 
en oe Le 
vy 3 nae he Of 

7 S| — _ te. 
: bony FEE, y “aryl | 2 


On Iridescent Plants 
by 


Davi W. LEE 


Botany Department 
University of Malaya, Kuala Lumpur 


Abstract 


Plants with leaves of iridescent blue and green colour are common in the deep shade 
of Malaysian rain-forests. Simple anatomical observations have revealed that the green 
iridescence is due to the refraction of diffuse light onto specially-oriented chloroplasts by 
lense-shaped cells. Blue iridescence colour is due to the operation of thin film interference 
filters in or on the epidermis. The advantage of such a filter in forest shade plants is the 
effective absorption of red wavelengths of light at the expense of the reflection of less 
important blue wavelengths. This report documents iridescence in the leaves of many 
unstudied taxa (mainly pteridophytes), common in Malaysian rain-forests. 


Life in a tropical rain-forest is not quite as easy for green plants as popular 
accounts would indicate. First by, the forest soils are generally rather sterile. 
Secondly, plants are under intense pressure of predation by insects and other 
animals, Finally, the light conditions are frequently less than optimum—from very 
bright conditions in the canopy to deep shade on the rain-forest floor. In this 
discussion we are concerned with the deficiency of light in the shade at the bottom 
of the rain-forest; in this environment can be seen a number of plants whose leaves 
display brilliant blue and green colours. These colours can be described as 
iridescent because of their intensity and metallic quality. The association of these 
iridescent plants with the shady and moist environments has compelled) many 
naturalists to reason that the colours must be associated with adaptations which 
help the plants to survive in the dark habitats. The rain-forest floor is one of the 
darkest environments in which plants grow. Not only is the total light quantity 
a small fraction of that above (less than one percent) but also the light quality is 
diminished; more of the wavelengths vital to photosynthesis, with the exception of 
some red light, have been absorbed by the foliage overhead (Evans, 1966; Federer 
and Tanner, 1966; Bjorkman, 1972). The purpose of this discussion is to explore 
how the iridescence of some Malaysian plants is associated with adaptations to 
increase the absorbance of diffuse and weak light for maximum photosynthetic 
efficiency. 


LEAF ANATOMY AND FUNCTION 


Leaves possess many structures which are significant in the efficient absorption 
of light in shady environments (see Ray, 1972; and Esau, 1965). Figure 1 sum- 
marizes these structures. 


The Cuticle. The function of the eit is two-fold (see Martin and J uniper 
1970 for further details). First, it repels water and prevents the rapid dessication 
of the leaf. Secondly, it can act as a selective filter in reflecting harmful ultraviolet 
light, and allowing the absorption of visible light essential for photosynthesis. 

The Epidermis. This superficial cell layer, particularly on the upper leaf 
surface, may possess special adaptations for efficient light absorption. It may have 
bumps which remain dry when the lower areas are covered by rainwater; these 
allow more efficient light absorption in wet conditions (Haberlandt, 1914). 


21 


22 Gardens’ Bulletin, Singapore — XXX (1977) 


Epidermal cells may also have regularly convexly curving upper surfaces; these 
focus diffuse sunlight onto specially orientated chloroplasts. Finally, the epidermal 
cells of extremely shade-tolerant plants (especially pteridophytes) may contain 
ee which can more directly receive light before it is scattered by the 
cell walls. 


Photosynthetic Parenchyma. Both the palisade and spongy mesophyll layers 
may have special adaptations for low light environments. They may be specially 
oriented beneath the lens-like epidermal cells. In shade-tolerant plants these cells 
would have fewer and larger chloroplasts. The protein/chlorophyll ratio, which 
reflects the relative plant investment in light reception versus biosynthetic 
apparatus, would be low. Finally, the cells may be closely packed together to 
minimize the reflection (and loss) of light between the cells. 


Stomata. In low light environments the stomata will not be a limiting factor, 
and we might expect various arrangements on the upper and, especially, lower 
leaf surfaces. 


Leaf Size and Shape. In low light environments leaves tend to be large and 
thin, with a high surface/volume ratio. This ratio is increased even more in plants, 
as ferns, with highly divided leaves. 


Thus, the task in understanding the significance of iridescence in shade-tolerant 
plants is to explain how some attributes of leaf structure confer an advantage on 
the plants and, at the same time, physically account for the iridescent colour. 


IRIDESCENCE 


Many plants from diverse taxonomic groups, from humid rain-forests 
throughout the tropics, possess iridescent leaves (Richards, 1952). Iridescent plants 
can be observed in Malaysian rain-forests (see Table 1), and some are extremely 
common. Selaginella willdenowii is a scandent pteridophyte frequently encountered 
in rain-forests, plantations and belukar throughout Southeast Asia. Another 
pteridophyte, Athyrium crenatoserratum, can be frequently observed in local rain- 
forest, as is Phyllagathis rotundifolia (Melastomataceae). Blue-leaved sedges of 
the genus Mapania commonly grow in Sarawak rain-forests, Others are less com- 
mon. The two kinds of iridescence observed in these plants, green and blue, are 
discussed separately below. 


Green Iridescence. The first observations on green iridescence were made 
on the cave moss, Schistostega, which grows in caves and other dark and wet 
environments in Europe and North America, Richards (1932) reported that the 
brilliant green colour encountered at particular angles was due to the functioning 
of certain protonemal cells as refractive lenses, focussing the diffuse light onto the 
specially oriented chloroplasts. Analysis of Malaysian green iridescent plants 
indicates that the same principles operate (see Figure 2 and Plate 1). These plants 
all share cells (epidermal or palisade) whose outer wails have a uniform convex 
curve. The chloroplasts are always oriented at the distal end of these cells. When 
we look at these plants the intense green colour can be observed only at an angle 
perpendicular to the surface of the plant. At other angles the colour is dull green 
or even whitish. This effect is due to the focusing of the diffuse light into a more 
intense beam directly onto the chloroplasts. These organelles, with their parallel 
stromal organization, absorb the light useful in photosynthesis and reflect the 
unused green light directly out of the long cells. Thus when we look at these plants, 
as at the entrance to Batu Caves near Kuala Lumpur or on rocks in especially shady 
forest habitats, we see the green light reflected in an intense and narrow beam. At 
other angles the light appearance of the leaves is due to reflection of whole light 
off of the cell surface acute angles and also the scattering of light within the cells. 
The birefringence of the cellulosic walls may also be a factor affecting the quality 
of the light reflected at acute angles. 


Iridescent Plants 23 


CUTICLE 
UPPER 
i a EPIDERMIS 
PALISADE 
AND MESOPHYLL 
PARENCHYMA 
, Oe cone ed 
AA e-\ 
J pel psn YT Ly eens VASCULAR 
+ Oee BUNDLE 
QO \ 


LOWER 
say —— EPIDERMIS 


Fig. 1. A diagram of the typical structure of a leaf. 


BRILLIANT GREEN COLOUR 
DULL GREEN COLOUR 
WHITISH YELLOW COLOUR 


LEAF SURFACE 


INTENSE GREEN 
LIGHT REFLECTED 


[TENSE GREEN 


DIFFUSE LIGHT FEIGHT REFLECTED 


BACK SCATTERING 
OF WHITE LIGHT 


DIFFUSE LIGHTK 
FROM EPIDERMAL | 
CELLS 


Fig. 2. The mechanism for the production of green light in iridescent green plants. 


24 Gardens’ Bulletin, Singapore — XXX (1977) 


Blue Iridescence. Men have been intrigued by the startling blue plants ever 
since they were first encountered. In the Malaysian rain-forest Selaginella willdenowii 
is the most striking example. In traditional languages the name ‘“‘peacock plant” 
is frequently used to describe these pteridophytes (as the Malay “‘paku merak’’). 
One aboriginal Senoi name for these plants translates roughly as “‘the hair on a 
tiger’s rump” (Gerard Diffloth, personal communication). These plants have been 
included in various medicinal remedies probably simply because of their colour. 
Selaginella willdenowii has been used in preparations for aching back as well as 
fever (Burkill, 1935). European scientists made frequent mention of the colour but 
gave no explanation. Stahl (1896) studied §. willdenowii while working in the 
famous Treub Laboratory at Bogor. He observed that the iridescent colour was 
due to pigmentation granules in the cuticle, Fox and Wells (1971) and Lee and 
Lowry (1975) recently published more adequate explanations for colour in 
Selaginella willdenowii, likely also to be the mechanism of colour formation for the 
other blue plants listed in Table 1. Stahl’s original observations can be discounted 
for three reasons. There are no presently known pigments in pteridophytes with 
the spectral properties to account for the blue colour. Secondly, in re-examining 
the epidermis of these plants we observed no pigment granules. Thirdly, the blue 
colour of the leaves disappears when they are covered with water. Thus, the colour 
must be due to some physical optical effect of the leaf surface, and not a pigment. 
There are three physical phenomena which can account for iridescent colour as 
seen in insects (see Michelson, 1911; Fox, 1959; Neville and Caveney, 1969). 
These could also be invoked to explain the blue leaf colour, Diffraction of light on 
thin gratings disperses different colours at different angles to the object’s surface. 
In S. willdenowii and the other plants, only blue colour can be observed. Tyndall 
scattering (which explains blue sky colour) polarizes light; no polarizing effects 
have yet been observed in any iridescent plants. Light interference, due to thin 
films, can produce uniform colours over relatively wide angles of incidence, and is 
the likely physical explanation of the phenomenon. 


The principle of a thin film (or quarter wavelength) interference filter is not 
familiar to most biologists, and requires some explanation. As a specific example, 
if the conditions described in Figure 3 are met, optical interference will result in 
the differential absorbance and reflection of different wavelengths of light. First, 
the filter must have a slightly different refractive index (r) than the surrounding 
medium, the air above or the cell below. Secondly, the filter must be exceedingly 
thin, a fraction of the wavelength of visible light. Thus, in the filter described in 
Figure 3, the thickness (142, ) is fraction of red visible light at 600 yp. If we trace 
the path of blue light (at 405 ») through this filter, we see that light of this wave- 
length will be reflected by the filter. The distance travelled from A and back to C 
equals the wavelength of that light in the filter. Retarding the light by one wave- 
length puts the light (visualized as a transverse wave) in phase, and the energy 
at this wavelength moves out of the filter. However, light with four times the wave- 
length of the filter thickness (or red light of 795 ») would be retarded half a wave- 
length at the same point, and its energy would be nullified. Thus such a filter can 
selectively enhance the reflection of certain wavelengths and allow the absorbance 
of other wavelengths. Two common examples are the rainbow colours seen from 
the thin oil films on rain puddles, and also the special anti-reflection coatings on 
quality cameras and binoculars. For a more technical explanation see Vasicek 
(1960). 


To test the above hypothesis Lee and Lowry (1975) analyzed the reflectance 
of light from both the iridescent and the older green leaves of S. willdenowii by 
using a slightly modified spectrophotometer. The analysis gave a peak of reflectance 
by the blue leaves at 405 p». They then constructed a filter model fitting this 
observation, as in Fgure 3. They assumed a filter refractive index slightly in excess 
of cell contents (of about r= 1.35), or about r= 1.4 as in Figure 3. In such a 
filter a concellation of the reflection effects would be observed at about 530 p, 


Iridescent Plants 25 


/ : AIR (r=1.00) 


FILTER (r=1-40) 


CELL (r=1-35) 


Fig. 3. Operation of a simple thin film interference filter. 


1. REFLECTS BLUE LIGHT. 


IN THE FILTER, AT AN EFFECTIVE WAVELENGTH OF 284, THE LIGHT IS 
DELAYED 4 WAVELENGTH AT POINT B AND IS NULLIFIED— CANCELLED AS 
OVERLAPPING TRANSVERSE WAVES WOULD SHOW. AT POINT C LIGHT IS 
DELAYED A FULL WAVELENGTH AND IS REFLECTED OUT — WAVES COINCIDE. 
AT THIS POINT THE REFLECTED LIGHT HAS A WAVELENGTH OF 4051 
(1.4/1.0 x 284) AND IS BLUE. 


2. NO EFFECT ON LIGHT AT 530 pu 
AT 2} THE FILTER THICKNESS THE WAVES WOULD CANCEL. 


3. ABSORBS RED LIGHT. 


IN THE FILTER LIGHT AT A WAVELENGTH of 568, IS DELAYED + WAVE- 


LENGTH AT C AND IS NULLIFIED— CANCELLED AS OVERLAPPING TRANS- 
VERSE WAVES WOULD SHOW. THUS MORE OF THIS LIGHT PASSES THROUGH 
THE FILTER AT B. HERE THE EFFECTIVE WAVELENGTH IN AIR IS 795 u 
(568 x 1.4/1.0) AND IS RED. 


with increased absorbance at higher wavelengths. In the difference spectrum 
obtained by subtracting values of the green leaves from the blue leaves; a curve 
similar to that predicted was observed (see Figure 4). Light above 700 » would 
be reflected by all leaves (Gaussman and Allan, 1973; Gates et al., 1965) and 
would obscure the filter effect at greater wavelengths. Thus, we have good physical 
evidence that such filters operate in the blue plants. 


The operation of such a filter has obvious adaptive significance for these 
shade-tolerant plants. Blue light (of which there is little in the dim light of the 
forest floor) is reflected, and the red light, which is more significant in the total 
photosynthesis by these plants, is absorbed. Thus, these filters can operate as 
“anti-reflection”? ccatings which increase the absorption of photosynthetically active 
light on the shady forest floor. 


Selaginella willdenowii plants possess additional adaptations to increase photo- 
synthetic efficiency. Firstly, the epidermal cells contain single large chloroplasts at 
the distal ends of the cells (see Plate 2). Thus, most of the photosynthesis occurs 


26 Gardens’ Bulletin, Singapore — XXX (1977) 


just below the leaf surface. Secondly, the epidermal cells have regularly convexly 
curved surfaces which appear to focus light onto the specially oriented chloroplasts, 
as previously described for the iridescent green plants, Finally, the iridescent leaves 
have protein/chlorophyll ratios significantly lower than those exposed to the sun 
(Krishnan, 1975). These plants possess a number of remarkable adaptations for 
the improvement of photosynthetic efficiency. The analogy of a camera with coated 
lenses and high speed film seems apt in describing the function of these plants. 


The other blue plants (see Table 1 and Plate 2) are different from Selaginella 
willdenowii in several details, although the same colour-producing principle 
probably operates for all of the plants. In S. willdenowii the blue colour can be 
removed by dipping the plants in water (blueness reappearing after drying). Other 
plants do not lose their blueness upon immersion. In the first case, the filter must 
be in some contact with the surface. In the second case the filter is likely to be 
found within the cell wall, and not directly exposed to the air. Scans of leaf 
reflectance for Athyrium crenatoserratum and Lindsaea scandens are similar to 
Selaginella willdenowii, except that the blue peak is at 410 p in Lindsaea and 
415 » in Athyrium. In both of the flowering species observed, Phyllagathis rotundi- 
folia and Begonia pavonina, I could not ascertain the wavelength of the blue colour 
because the colour was effectively produced when the incoming light was at a 
smaller angle to the surface than the reflected light. The method of measurement 
required a fixed incident angle of 60° for both. The physical constitution of the 
filters is not presently known, but in all cases the colour is due to a physical optical 
effect and not to a pigment. 


Other blue plants possess distinct anatomical features in addition to Selaginella 
willdenowit. In Athyrium crenatoserratum (Plate 2), the epidermal cells also 
contain chloroplasts, and the outer wall has the convex curve described above. The 
blue colour is particularly apparent in the older leaves of this common fern, Perhaps 
the most spectacular iridescent blue plant of all is the rare begonia, Begonia 
pavonina. When looking directly at these plants one sees an intense dark blue 
colour. However, if the angle of observation is changed so that light is reflected 
at a more acute angle, the colour changes from blue to dull green and then to gold. 
An interference filter would account for the blue colour, and the gold colour is 
probably due to the refractive properties of the epidermal cells. The plant is now 
under further study. The chloroplasts of Begonia pavonina are at the bottom of the 
palisade parenchyma cells. Both light refraction and interference appear to enhance 
light absorption in this plant, as in Selaginella. The palisade cells are packed very 
closely together to minimize light loss due to scattering effects. Immediately beneath 
is a compact mesophyll layer, the cytoplasms of which is dark red with antho- 
cyanins. This layer probably reflects red light (comparatively more abundant on 
the rain-forest floor) right back up to the chloroplasts in the palisade cells. Thus 
the Begonia pavonina plants appear to have leaves with lenses, filters and special 
mirrors—all to increase conversion of precious light energy to sugars in the shady 
forest environment. 


FURTHER RESEARCH 


Rain-forest floor plants, as in Malaysia, have some spectacular adaptations for 
the efficient utilization of light. Further studies may uncover new principles, and 
will help illuminate the functional significance of anatomical details of leaf 
structure. I see some specific areas of research potential. 

First, although there is now some evidence for the operation of thin film 
interference filters in plants, no one has yet been able to elucidate the structural 
basis of the filter in the different plants. Lee and Lowry (1975) discounted the 


t 


‘ 


Iridescent Plants 27 


9 

2) 

c 

= 

1S) 

s 

2 

wd 

= 

5 

ST] 

a 

500 
Wavelength / 

Fig. 4. The relative reflectance of light by green (— — — — — — ant ie 
Selaginella leaves as recorded by a spectrophotometer. The bottom curve (——:————) 


is the difference between the two curves above and gives a plot similar to that expected for 
a thin film interference filter. 


cuticle as being the molecular basis of the filter in S. willdenowii; non-polar 
solvents, which dissolve cuticle waxes, did not remove the interference. Fox and 
Wells (1971) speculated that the structural basis for the optical effect may be the 
trapping of thin layers of air by the epidermis, as is well known for some insects. 
Critical experimental support for the optical phenomenon of interference should 
include physical information on the nature—including the thickness and refractive 
index—of the filter. The small dimensions will necessitate the use of sophisticated 
electron microscope techniques. If the thickness and refractive index of the filters 
were known, then we could precisely predict the nature of the interference effect. 
The basis for the filter may well lie in the orientation of cellulose microfibrils in 
the cell wall; further research will solve this problem. 


Rather esoteric studies on the leaves of shade-tolerant plants may eventually 
be of some practical importance, In designing more efficient plantation agriculture 
systems for the tropics, biologists will have to consider the kinds of plants to grow 
in the understoreys of the plantations. Knowledge of leaf structure may help us 
select plants which could thrive in such environments. Who knows? maybe 
someday we will spray interference filters onto the leaf surface of understorey 
plants, making them blue—and more productive. 


Acknowledgements 


The following individuals were extremely helpful in the various stages of the 
preparation of this manuscript. Prof. R. E. Holttum gave encouragement and 
additions to Table 1; Mr. Willis Littke also gave additions. Drs. B. C. Stone, 
J. B. Lowry and Peter Ashton, and Prof. Paul Richards gave advice and encourage- 
ment. Prof. Anne Johnson identified the liverwort. Encik Jayamohan Krishnan and 
Encik Ho Kok Wai provided photographs; Cik Tan Geok San gave technical help 
and Cik Wan Shahrizah typed the manuscript. 


28 Gardens’ Bulletin, Singapore — XXX (1977) 
References 


Bjorkman, O. 1972. Photosynthetic adaptation to contrasting light climates. 
Carnegie Institution Year Book 71: 82-135. 


Burkill, I. H. 1935. A dictionary of the economic products of the Malay Peninsula, 
two vols. London, Crown Agents for the Colonies. 


Esau, Katherine. 1965, Plant anatomy, second edition. John Wiley and Sons, 
New York. 


Evans, G. C. 1966. Model and measurement in the study of woodland light 
climates. In R. Bainbridge, G. C. Evans and O. Rackham, eds. Light as an 
ecological factor. Blackwell Scientific, Oxford. 


Federer, C. A. and L. B. Tanner, 1966. Spectral distribution of light in the forest. 
Ecol. 47: 555-560. 


Fox, D. L. 1953. Animal biochromes and structural colours, Cambridge University 
Press, London. 


—_——-—— and James R. Wells. 1971. Schemochromic blue-leaf surfaces of 
Selaginella. Amer. Fern. Jour. 61: 137-139. 


Gates, D. W., H. J. Keegan, J. C. Schleter and V. R. Weidner. 1965. Spectral 
properties of plants. Applied optics 4: 11-20. 


Gausman, H. W. and W. A. Allen. 1973. Optical parameters of leaves of 50 plant 
species. Plant Physiol. 52: 57-62. 


Haberlandt, G. 1914, Physiological plant anatomy. MacMillan and Company, 
London. 


Krishnan, Jayamohan. 1975. Physiological leaf anatomy of Selaginella. B.Sc. 
Honours Thesis in Botany, Faculty of Science, University of Malaya, Kuala 
Lumpur, 


Lee, D. W. and J. B. Lowry. 1975. Physical basis and ecological significance: of 
iridescence in blue plants. Nature 254: 50-51. 


Martin, J. T, and B. E. Juniper. 1970. The cuticles of plants. Edward Arnold, 
London. 


Michelson, A. A. 1911. On metallic. colouring in birds and insects. Phil. ae 
21: 554-567. 


Neville, A. C. and S. Caveney. 1969, Scarabaeoid beetle exocuticle as an oma 
analogue of cholesteric liquid crystals. Biol. Rev. 44: 531-562. 


Ray, Peter, M. 1972. The living plant. Holt, Rinehart and Winston, New York. R 


Richards, P. W. 1932. The ecology of bryophytes. In Fr. Verdoorn, ed:: Manual 
of bryology. Martinus Nijhoff, The Hague. 


1952. The tropical rainforest. Cambridge University Press, London. 


Stahl, E. 1896. Uber Bunte Labblatter. Ein Beitrag zur Pilanzenbiolage II, Ann. 
Jard. Bot. Buitenzorg 13: 137-216. ‘“ 


Vasicek, A. 1960. Optics of thin films. North Holland por pte: Company, 
Amsterdam. 


ON 
N 


Iridescent Plants 


"SVoIe UIVJUNOW UI SUOTZVAD[O WINIPOU 3e 


"YEMVICS 


uowwo,) 


ul sjuvjd onjq  ywodsoprit 


‘poutn} si 
Jeo] 94} SB plod 0} BUIUIN} SIq} ‘soAvOT 
pommoyjoo ysidind-ysinjq 94} IOJ SUTyINYS 


‘yuejd yuddsopldI on]{q 
UOWWOS SOU 94} MMOUAap]IM *S IOV 


edoD (Ia) wnyofips09 
wnucyjp 10} poylodsl oOs[ty ‘usoys 
onjq o[eJouU e OARY SOALIT JOP[O 34] 


"SOAVOT SY} 0} UDOYS 
o[ejour onjq e& savy AewW (‘[q MpiION 
"TT sdeyiod) vapspury jo soisods I0yIO 


“Bulidgs (TI@A\) 
SIPIMIAOLD “S SY ‘yUSDSOpIIt 9q ose 
Aewut soiods IoyJQ ‘jur[d poavoj-on[q 
porsjunosus ApUoNboJJ Jsow oy} sdeylog 


"JU9}Xx9 
IISso] B O} jNQ ‘SoAvdT U99I3 JUSOSOPLII 
sary Os[e soldods nIuosag I9YIO *jSO1OJ 
ul syoo1 dwep pue Apeys uo uoWWOoD 


"JUD}X9 IOsso] & 0} JNQ ‘JUSOSOpLIT 
St IPA Npjaysioy “W poyeel oyL 
‘SOARVDZ) NYVG SAvd AlIeNjOURS SPISUI Ud9¢g 
‘ud013 yUDOSOpIM Ivodde suojeoso] yep 
AIOA Ul MOIS yey} Xe} IOYJO pue si} 
‘SOARD Nyeg ‘oAvd Yep JO YNOU je UD9G 


SLNHNWOO 


‘eIsy yseo YNOg UI sIOYM 
-osjo Ssdeyiod puve ‘viske[eyy ISOM 


‘eIsy yseq yNOS 
UOI}JLAT[I YSIY O} MOT WOIT 


ysol0y dooq ‘visAeleyy JSOAA Ul DIY 


‘eISY ySeq YyINOS 
UI }SOIOJ UOVA[D MOT UI UOWWOD 


“VISAR[RIY 
SOA, ‘SAOTTVA UleyUNOW YsIY UT 


"eISY “A ‘S Ul 4solOJ 
UONVAId-pru puke MOT UI UOWUWOD 


‘ejnsuluod oy} nNoYysnoIY} 
sAg][eA UleJUNOW pue 4saIOJ MOT 


‘sosnoy 
-u9013 «= oyelodwia} ~Ul os[e pue 
vIsy “q ‘§ Ul UOWUWOS AjoWoIyxA 


SLINV1Id AN 1d 


eIsy “d 'S 


"eISAR[RIL “AA Ul 
sdo19jno SUO}SOWI] Moj & O} POPTWIT 


BIsy “Ad 'S 


NOILNETALSIG 


(svoprlopiio}doAiq) 
9BvIORI}pPoRSUUICG] 


avoovIodAy 


ovooRIUOsog 
IVIOLJVUIO}SL[I 
dvd0eRI}pov}suusq 


(avaprloplis}doAIq) 


(svoplolAyiy) 
avoovl}por}suusg 


(avoplooespurT) 
oevddRI}poV}suudq, 


DVIOL|[OUIB IIS 


aeoovIUOsog 


OBOOBUTWTLS|[e 


(episdosedoayy) 
ovooRURYyoOIe|] 


ATINVA 


"ws ‘¢ (MS) DpNJDOUNLT 
Duapj]yIoulcpiq 


‘ds viundvpy 


[Ply vuluoand viuosag 
Ta Pyofipunjos siyiwsn]j<yd 


‘WIOH CHIOH) Wiuny sisdoyouskjog 
OPIN (Ig) wnivssasojpuasia wniudyip 


‘WOH Slajsassa] “IeA 
‘YH Suapuvos vavspurT 


‘yeq (Aso) NMouappyim vjaursvjag 


‘TIE DIDNUIS DIUOsag 


[Pry suajnod vanjjkydouop 


‘uyYyoS wnuassipyao{ unipoyjwky 


NOXVL 


ea ae tee eS ee 


SLNVTId NAdaaXD 


RISAR[VJL Ul Poarosgo syuR[d yUSOSOpIII JO 3s!I] V 
. NN _eeeEeEeEeEee EEE 


CLAD 


Mioaq2 Sines eitt0 Jew} | 
a 5 aetahii : ae 
seeire ~~ ae 7 


yori teosa 


acbC Jnaig 5 7 ‘ > ¢ + j 
» « ’ . an A 4 
mit ee 218: : ks : 4 . -m 2% 
. ° ~ - 


Bai Mu ve Myer abies 
+ Sia tee , : ; 2 mk 


sHotd (tome ® Ein wager 


48 Se eget a8 


= - 
somthes ior onl. ierowesls\\\ | 


ae Beinbla>  Azilqreg-riuic 
. Yesl or && bios Gi : 


c 
m .@igniq- ‘suid_- tas 


-— . 
eee Hicthvony th eno 


Plate 1. Photographic details of the leaf anatomy of green iridescent plants. A — Transverse 
section of a Cyathodium foetidissimum thallus showing the transversely curved epidermal 
cells and the distally oriented chloroplasts. B— Upper leaf surface of Monophyllaea patens 


showing the long and thick-walled palisade cells which help deflect the green light in a 
narrow path out of the leaf. 


Plate 2. Photographic details of the leaf anatomy of blue iridescent plants. A — Transverse 
section of Selaginella willdenowii leaf. Note the peculiar shape of the epidermal cells and the 
large single chloroplasts. B— Scanning electron micrograph of the surface of the S. will- 
denowii leaf; the convex curvature of the epidermal cells is easily seen. C — Transverse 
section of leaf of Athyrium crenatoserratum. Note the convexly curving epidermal cells. The 
two transverse-section photographs are courtesy of Encik Jayamohan Krishnan; SEM micro- 
graph was supplied by Encik Ho Kok Wai. 


The Ferns of Gunong Ulu Kali 
by 


A. G. PiccoTT * 


Summary 


The various habitats on the peak of Ulu Kali and in the surrounding area, the Genting 
Highlands, are described. Some one hundred species and varieties of ferns found between 
5,000 feet and the summit are listed. 


Gunong Ulu Kali is the southernmost high mountain peak in the Main Range 
of Malaya, and is only 20 miles NNE of Kuala Lumpur. The environment and 
vegetation of the mountain and surrounding area, the Genting Highlands, have 
been described briefly by Burgess (1969). Before the Hotel and Casino Complex 
was opened in 1971, this part of the Main Range could only be reached by walking 
and climbing through the jungle. Now, at Genting Simpah, a road branches off the 
main Kuala Lumpur/Kuantan trunk road and winds its way to the top. At about 
5,000 feet, with one mile still to go, the character of the vegetation changes. The 
trees are smaller, gnarled and mossy. There are fewer climbers, abundant lower 
down, and fewer tree ferns. Higher still, the vegetation changes again and becomes 
the dwarf and scrub forest of the Cloud Zone. The trees here are stunted; there is 
little or no grass and epiphytes are abundant. 


Though few people venture beyond the Complex, the road continues. Your 
vehicle can take you within yards of the summit of Gunong Ulu Kali (5,814 feet), 
and along the whole length of the ridge to Bukit Genting Chin Chin. The slopes 
are steep or precipitous, and treacherous. On a clear morning the views from the 
ridge are spectacular: neighbouring mountain peaks covered with jungle, the City 
of Kuala Lumpur, the tin mines, and, in the far distance, the sea. It is cool, 
refreshing and still. If cloud descends, it becomes chilly, visibility is limited to a few 
yards and strong winds may blow. 


The area studied includes Gunong Ulu Kali and the associated ridge, and also 
some lower slopes towards the two pumps supplying the Complex with water. The 
ferns listed were all found between the summit and about 5,000 feet. It was not 
possible to record every species in all places, but the list does give an indication of 
their frequency and distribution. For convenience, the area was divided into a 
number of locations, each the centre of forays into the various habitats nearby. 


Ferns of Malaya (Holttum 1966) was used as an aid to identification, 
but since the publication of the second edition, the author has reappraised the family 
Thelypteridaceae (1971) in which sense the generic names are used here, Specimens 
were deposited at the Herbarium of the Royal Botanic Gardens at Kew, and an 
incomplete set also at the Botanic Gardens, Singapore. 


DESCRIPTION OF LOCATIONS 


A is situated at the end of the road along the ridge, 1.75 miles from the car 
park. The Radar Station occupies the small peak just beyond. There are still 
remnants of scrub forest nearby, and lower down the steep slopes is montane 
ericaceous forest, with conifers and Pandanus. Earthmoving has been carried out 
on a large scale, leaving cleared areas and fresh earth banks. Drainage is channelled 


*21, Jalan Dato Klana, Seremban, Malaysia. 


31 


ae Gardens’ Bulletin, Singapore — XXX (1977) 


into a steep mossy valley on the eastern side, and this was where Habenaria 
angustata, a terrestrial orchid, was found. It was growing in mossy peat and on 
rotting tree-trunks. 


B is 0.15 mile along the road, back towards the Hotel Complex. The ground 
drops away sharply to the east. On the opposite side of the road are wet mossy 
hollows with Pandanus klossii, and above them dwarf forest. The trees support an 
abundant flora of mosses and lichens, and epiphytic ferns and orchids. In September 
and October some trees were almost covered with pinkish purple orchid flowers. A 
large crested lizard was first seen here. 


Equisetum debile was growing in sand near the roadside drain at 0.25 mile. 
Over a period of one and a half years it developed from a small tuft to a large 
straggling mass. 


C is 0.3 mile along, at the point where the road changes from one side of the 
ridge to the other, Strong winds blow through the gap. Large granite boulders form 
pinnacles with tree-roots growing over the weathering surfaces. Above the road is 
dwarf mossy forest; at road level there are cleared areas and earth banks; and then 
the ground drops away on either side. Pholidota parviflora grew here, and Coelogyne 
sp. with fleshy red scapes and drooping racemes of flowers on long pedicels. 
Further on, the road has been blasted out of the rock of the ridge and there is 
a precipitous drop of about 1,000 feet on the western side. Here, an almost 
horizontal vein — apparently decomposing felspar — has been exposed. The road 
then crosses back to the east side of the ridge. 


D, at 0.4 mile, is rather open. In the valley are larger trees, raising their crowns 
to the level of the ridge, and tall tree ferns. Earth banks border the road. Fragrant 
Coelogyne longibracteata and Dendrochilum simile were common and flowered 
towards the end of the year. 


E is at 0.5 mile, with large trees in the valley below. The stunted forest above 
contains a high proportion of conifers which have the habit of dropping the ends 
of their branches. Bulbophyllum angustifolium was in flower in February. There 
are wet mossy hollows where epiphytic ferns are abundant and the fruiting bodies 
of macroscopic fungi can be found. 


F is very exposed at 0.8 mile, the road running almost along the crest of the 
ridge. There are fresh earth banks, patches of secondary growth, fallen trees, dwarf 
forest and the heads of small steep valleys. An established path goes into the forest 
behind but care should be taken: it often becomes a running stream, and there 
are scorpions. 


G, 0.95 mile, is somewhat sheltered. The trees are larger and some have fallen, 
making identification of epiphytes growing in the crown much easier. Coelogyne 
longibracteata was abundant and flowered late in the year, Dendrobium cornutum 
and D. atrorubens produced flowers in July. Wild bananas, ginger and a variety of 


creepers can be seen. Terrestrial ferns with large fronds grow on the edge of the 
clearing. 


H is 1.1 miles along the ridge and there is a large clearing, an abandoned 
building site, to the east. Beyond this a path leads into scrub forest on a short 
northerly ridge. The stunted trees, which included Dacrydium beccarii and Rhodo- 
dendron orion, are barely 6 feet tall and grow in thick peat and sphagnum root 
mat. An arborescent grass was growing near the end of this ridge. Dischidia 
astephana and some Nepenthes spp. were also here. In July many flowers of 
Habernaria angustata were found. Dwarf forest grows on the other side of the road 
and mossy hollows drain into streams which pass under the road. 


1: Gunong Ulu Kali, 1.25 miles along the ridge and 0.5 mile from the Complex. 
The Television Station has been built on the summit, but just below and across the 
site of the construction workers’ hut is an area of nearly undisturbed scrub forest. 


Ferns of Ulu Kali 33 


ery ; 
niin 
Chin Chun 


GENTING HIGHLANDS 


V4 V2 


miles 


V726 


goo? 


TV} 
Guyer = 


ae (1) oS 5 
Mushroome 
Farm} 


Western 
Dam 5: ) 
\* R 
\: Cable Car 
yy ’ : 
4; i : : 4 
fi! ores Bork 4 
4, oa . 


* a 


Incinerato 
be ~ 
“S 
> 
> 
TS" % 
Sy 
7eXo> 
4 
$5 


KEY: 
otorable roc? = ====m=== ay fo, 
Jeep track — 
Track 


The twisted trees are covered with mosses, lichens and epiphytes. From here the 
road winds downhill, with steep debris-covered slopes to the east. Further still the 
road has been cut into the side of the mountain and the natural environment has 
been much disturbed. Little remains of the natural vegetation: pioneer species are 
becoming established. 


34 Gardens’ Bulletin, Singapore — XXX (1977) 


Locations J, K and L cover larger areas, down to an altitude of about 5,000 
feet: 


J is the land adjacent to the track which leads to the eastern dam and pump 
house, and begins north of the main hotel and beyond the boating lake. It descends 
some 1,000 feet, running round the mountain-sides. Deep road cuttings have been 
made in the steep slopes. The incinerator and main refuse dump are situated along 
here. This area is much more sheltered than the ridge and some very large trees 
grow in the main valleys. Near the dam there are small swamps which drain into 
the streams. 


K begins just beyond the incinerator, where a rough track bears off to the 
right towards Gunong Lari Tembakau. For a while a pack of ‘wild’ dogs inhabited 
those parts of it where the ground has caved in. Dwarf and scrub forest clothes 
the peaks, but the slopes are shaded by taller trees. There are some large mossy 
boulders in one of the wetter patches, and amongst these Cryptostylis arachnites 
and Ceratostylis ampullacea were flowering in January. 


L is along a track which turns off the main road, one mile down and just below 
the residence ‘Sri Genting’. It more or less follows the 5,000 foot contour and leads 
to the western dam and pump-house, directly under the Television Station and the 
Mushroom Farm, The track has been cut into the side of the mountain. It passes 
through fairly tall sheltered forest, crossing several streams. Wet rock faces are 
found near the streams, in places covered with mosses and liverworts, including 
Marchantia sp., but few Hymenophyllaceae. 


LIST OF GENERA 


Ferns occurring on Gunong Ulu Kali, above 5,000 feet, arranged in families. 


Schizaeaceae Grammitidaceae Oleandroideae subfam, 
Schizaea Grammitis Nephrolepis 
Matoniaceae Bae ey Oleandra 
; alymmodon nae 
Matonia M : Pteridioideae subfam. 
Ctenopteris hs 
Gleicheniaceae Scleroglossum Pieris | 
Gleichenia Histiopteris 
Dicranopteris Thelypteridaceae Pteris 
Hymenophyllaceae ee 3 bide Asplenioideae subfam. 
ingia 
Hymenophyllum ! ; Asplenium 
Cory phopteris 
Trichomanes S phaeroste phanos Blechnoideae subfam. 
Plagiogyriaceae Pneumatopteris Blechnum 
Flagiogyria Christella Lomariopsidoideae subfam. 
Cyatheaceae Dennstaedtiaceae Elaphoglossum 
Cyathea Dennstaedtioideae subfam. Teratophyllum 
Polypodiaceae Microlepia Dryopteridoideae subfam. 
Dipteris Hy polepis Acrophorus 
Belvisia 
Li ‘ Ms airy 
Gaines indesepidene subfam. Athyrioideae subfam. 
; - Lindsaea Diplazium 
silica § phenomeris 
Cry psinus ss Adiantaceae 
Goniophlebium Davallioideae subfam. Pityrogramma 


Lecanopteris Davallia Vittaria 


Ferns of Ulu Kali 35 
List OF SPECIES AND VARIETIES 


Ferns occurring on Gunong Ulu Kali, above 5,000 feet, arranged alphabetically. 
with locations and brief notes. 

1. Acrophorus blumei Ching apud C. Chr. Plate 1. 
Becations:,:AlbEe-G, K:. L. 
In shady hollows and valleys. 

2. Asplenium caudatum Forst. 
Locations: J, K, L. 
On rocks by streams. 

3. Asplenium nidus Linn. 
Locations: J, L. 
Epiphytic on larger trees in the deeper valleys. 

4. Asplenium scortechinii Bedd. 
Location: J. 
Epiphytes on mossy trees near the dam. 

5. Asplenium unilaterale Lam. 
Location: L. 
In rocky stream-bed. 

6. Belvisia revoluta (Bl.) Copel. 
Locations: J, L. 
Epiphyte on mossy trees in wet hollows. 

7. Blechnum orientale L. 
Locations: H, and between I and Hotel Complex. 
On earth banks and exposed rock faces, often stunted. 

8. Blechnum vestitum (Bl.) Kuhn 
Locations: A, B, C, E, I J, K. 
In dwarf forest with Plagiogyria tuberculata. 

9. Calymmodon cucullatus (Nees & Bl.) Pres] 
Location: J. 
Small epiphyte on mossy trees near dam. 

10. Chingia pseudoferox Holtt. 
Locations: I, L. 
In the open by roadside drains. 

11. Christella arida (Don) Holtt. 
Location: I, 
In open by roadside drain. 

12. Coryphopteris badia (v.A.v.R.) Holtt. 


Location: G., 
First record for Malaya. 
In light shade in mossy hollows. 


13. Coryphopteris gymnopoda (Bak.) Holtt. 


Locations: A, B, E, I, L. 
In light shade in dwarf forest. 


36 


14. 


., 


iT. 


18. 


20. 


ZA; 


pHs 


23s 


24. 


25. 


26. 


ar. 


Gardens’ Bulletin, Singapore — XXX (1977) 


Coryphopteris gymnopoda (Bak.) Holtt. var. bintangensis Holtt. 
Location: B. 

On forest edge. 

Crypsinus enervis (Cav.) Copel. 

Locations: A, D, E, G, H, J, K. 

Epiphyte in moderately exposed places. 
Crypsinus laciniatus (Presl) Holtt. Plate 2. 
Locations: B, E, G, H, J, K, L. 

Epiphyte in open places. 

Crypsinus wrayi (Bak.) Copel. 

Locations: A-C, E, G-L. 

Small epiphyte on very mossy trees. 
Ctenopteris contigua (Forst.) Holtt. Plate 2. 
Location: H. 

Epiphyte in mossy hollow. 

Ctenopteris fuscata (Bl.) Kze 

Locations: E, J. 

Small epiphyte in dwarf forest. 

Ctenopteris khasyana (Hk.) Holtt. 

Location: E, 

Epiphyte in mossy hollow. 

Ctenopteris mollicoma (Nees & Bl.) Kze 
Locations: A, J. 

Epiphyte in exposed situations. 

Ctenopteris moultoniit (Copel.) C. Chr. & Tard. 
Location: J. 

Epiphyte in sheltered valley. 

Ctenopteris obliquata Copel. 

Location: J. 

Epiphyte in sheltered valley. 

Ctenopteris tenuisecta (Bl.) J. Sm. Plate 1. 
Locations: H, J, K. 

Larger epiphyte in deep valleys. 

Cyathea contaminans (Wall. ex Hook.) Copel. 
Locations: E, G, I, J, K. 

Large tree fern, on edge of forest and in valleys. 
Cyathea hymenodes Mett. 

Locations: F, G, H, J. 

Tree fern on edge of dwarf forest. 

Cyathea hymenodes Mett. (variety). Plate 1. 


Location: H. 

Tree fern on edge of clearing. I consider this sufficiently different from the 
species in its much-reduced pinnae on the base of the stipe, to call it a variety 
for the time being. 

Collections. 21.06.75: 1334 (SING), 1335, & 1337 (K), 1338 (SING). 


Ferns of Ulu Kali 37 


28. 


29. 


30. 


31, 


ad. 


a3, 


34. 


a0. 


36. 


af. 


38. 


39. 


40. 


Cyathea lurida (Bl.) Copel. 
Locations: B, C, E, F, G. 
Tree fern in dwarf forest along ridge. 
Davallia trichomanoides BI. 


Location: J. 
Epiphyte in sheltered valley. 


Davallia trichomanoides Bl. var. lorrainii (Hance) Holtt. 
Location: L. 

Epiphyte, in light shade. 

Dicranopteris curranii Copel. 


Locations: F, G, J, K. 
On fairly exposed earth banks. 


Dicranopteris linearis (Burm.) Underwood var. linearis Holtt. 


Locations: B, H, I. 
Colonising exposed clearings. 


Dicranopteris linearis (Burm.) Underwood var. montana Holtt. 


Locations: F, H, J. 
On side of road cuttings. 


Dicranopteris pubigera (Bl.) Nakai 


Locations: G, I. 
On roadside banks in exposed situations. 


Diplazium accedens (B1.) Milde 


Location: L. 
In small sheltered valley. 


Diplazium asperum Bl. 


Locations: E, G, H, J, L. 
On edge of forest. 


Diplazium speciosum Bl. 


Locations: B, C, D. 
In fairly exposed situations near end of ridge. 


Dipteris conjugata Reinw. 
Locations: G, H, I, J, K. 
Colonising steep earth banks. 


Elaphoglossum sp. Plate 2. 

Locations: A, B, C, D, F, G, H, J, K. 

Epiphyte in less exposed places. 

A new species which will be published and probably as E. robinsonii. 
Differs from E. callifolium and E. malayense in having pale brown, thin, flat, 
entire scales and very broadly pointed lamina. It matches exactly an 
incomplete specimen collected by H. C. Robinson in 1913 on _ nearby 
Gunong Mengkuang. 

Collections. 12.10.74: 1093 (K); 9.11.74: 1095 (SING); 10.11.74: 1094 (K); 
10.08.75: 1339 & 1340, 1344-1347, 1358 & 1359 (all K). 


Gleichenia hirta Bl, var. amoena (v.A.v.R.) Holtt. 
Locations: F, G, I. 
In fairly exposed grassy clearings. 


38 


41. 


42. 


43. 


44. 


45. 


46. 


47. 


48. 


49. 


50. 


>i. 


ae 


a3 


54. 


Gardens’ Bulletin, Singapore — XXX (1977) 


Gleichenia longissima Bl. 
Locations: F, G, H, I, J. 
Scrambling on edge of forest. 
Gleichenia microphylla R. Br. 


Location: I. 
In scrub forest and grassy clearing. 


Gleichenia truncata (Willd.) Spr. var. plumaeformis (Presl) Holtt. 
Locations: B, G, 1, .L. 

Scrambling on edge of forest. 

Gleichenia vulcanica Bl. 

Locations: F, G, H, I. 

In and on edges of dwarf and scrub forest. 
Goniophlebium persicifolium (Desv.) Presl 
Locations: J, K. L. 

Epiphyte in sheltered places. 
Goniophlebium prainii (Bedd.) C. Chr. 
Locations: E, G, L. 

Epiphyte on larger trees in valleys. 
Grammitis hirtella (Bl.) Tuyama 
Locations: “HAI 9: 

Epiphyte in dwarf and scrub forest. 
Grammitis hirtella (B\l.) Tuyama var. major Holtt. 
Locations: A, H, I. 

Epiphyte in very mossy dwarf forest. 
Grammitis reinwardtii BI. 

Locations: A, B. 

Epiphyte on mossy trees. 

Histiopteris incisa (Thunb.) J. Sm. 
Locations: D, I, Fes. 

Scrambling, on edge of forest. 

Histiopteris stipulacea (Hk.) Copel. 


Location: K. 
On edge of small clearing. 


Hypolepis punctata (Thunb.) Mett. 

Location: E. 

Few patches of small fronds in sandy clearing. 
Hymenophyllum acanthoides (v.d.B.) Rosenst. 
Locations: A, E, G. 

Abundant on some mossy tree-trunks. 
Hymenophyllum blandum Racib. 


Locations: B, H. 
Small epiphyte in dwarf forest. 


Ferns of Ulu Kali 39 


55. 


56. 


awe 


58. 


59. 


60. 


61. 


62. 


63. 


64. 


65. 


66. 


67. 


68. 


Hymenophyllum exsertum Wall. ex Hook. 
iLocauoes:) A. B, C, H, LK. L. 
Epiphyte on mossy trees. 
Hymenophyllum javanicum Spr. 


Location: J. 
Epiphyte in valley near dam. 


Hymenophyllum serrulatum (Presl) C. Chr. 


Locations: I, K, L. 
Epiphyte in less-exposed forest. 


Lecanopteris carnosa (Reinw.) BI. Plate 1. 

Locations: H, K, L. 

Epiphyte in crowns of trees. Easily accessible in scrub forest of small ridge. 
Lindsaea malayensis Holtt. 

Locations: E, G, H, J, K. 

In wet mossy rocky hollows, and sometimes on rotting fallen trees. 
Lindsaea oblanceolata v.A.v.R. 

Locations: H, K, L. 

Climbing in wet hollows; L. pectinata of ‘Ferns of Malaya’. 

Lindsaea rigida J, Sm. 


Location: I. 
In sheltered parts of mossy forest, climbing. 


Loxogramme avenia (Bl.) Presl 


Location: J. 
Epiphyte and on rocks in wet valley near dam. 


Macrothelypteris torresiana (Gaud.) Ching 
Locations: A, I. 
In the open by roadside drains. 


Matonia pectinata R. Br. 
Location: I. 
On the top of roadside bank, very exposed, near summit. 


Microsorium sarawakense (Bak.) Holtt. 
Location: L. 
Epiphyte near dam. 


Nephrolepis davallioides (Sw.) Kze 
Locations: E, J. 
Epiphyte on few large trees in deeper valleys. 


Nephrolepis tuberosa (Bory) Presl 
Locations: I, J. 
Fronds small, in peat at edges of clearings. 


Oleandra pistillaris (Sw.) C. Chr. 

Locations: G, H, J, K, L. 

Straggly shrub in open sheltered places, Plants of a different habit, creeping 
and producing fronds singly, found on the middle part of the ridge. 


69. 


70. 


Th. 


Fa. 


73. 


74. 


75. 


76. 


77. 


78. 


79. 


80. 


81. 


Gardens’ Bulletin, Singapore — XXX (1977) 


Pityrogramma calomelanos (L.) Link 

Locations: B, E, I, J, K. 

Usually a small plant, rapidly becoming established on earth banks and in 
clearings. 

Plagiogyria tuberculata Copel. 

Locations: A, B, C, E, F, J, K, L. 

Abundant in wet dwarf forest, with Blechnum vestitum. 

Pneumatopteris ecallosa (Holtt.) Holtt. 


Location: G. 
In shade on side of valley. 


Pteridium aquilinum (L.) Kuhn var. wightianum (Ag.) Tryon 
Locations: D H, I. 


Colonising earth banks and small clearings, and rapidly becoming more 
common. 


Pteridium caudatum (L.) Maxon var. yarrabense Domin 


Location: H. 
One patch on edge of clearing. 


Pteris longipinnula Wall. 


Location: 1... 
In shelter of forest. 


Pteris tripartita Sw. 


Location: I. 
Single well-developed fertile plant on steep slope of debris. 


Pteris vittata L. 


Locations: B, E, I. 
Common by side of road, but fronds often small. 


Schizaea malaccana Bak. 


Locations: H, I. 
In moss on dwarf trees in sheltered hollows. 


Scleroglossum minus (Fée) C. Chr. 
Locations: A, B, G, H, I, J, L. 
Epiphyte in mossy forest. 


Scleroglossum pusillum (Bl.) v.A.v.R. 
Location: H. 

Epiphyte in scrub forest on small ridge. 
Sphaerostephanos polycarpus (Bl.) Holtt. 


Locations: A, B, I. 

In exposed situations by roadside drains, Differs from lowland specimens in 
having a more hairy lower surface and lacking glands on the upper surface, 
but matches one collected on Taiping Hills by Day in the 1880s. 


Sphenomeris chinensis (L.) Maxon var. divaricata (Chr.) Kramer 


Locations: G, J, L. 
On steep earth banks and cuttings. 


Ferns of Ulu Kali 41 


82. Teratophyllum aculeatum (BI.) Mett. var. montanum Holtt. 
Location: L. 1} 
High-climbing, clothing trunks of tall trees in valleys; upper limit of species 
about 5,000 feet. 
83. Trichomanes meifolium Bory 
Locations: A, B, E, F, H, I. 
Epiphyte in mossy dwarf forest. 
84. Trichomanes obscurum Bl. 
Locations: C, J. 
In wet peaty hollows. 
85. Trichomanes pallidum BI. 
Locations: G, H. 
Epiphyte on shadier side of mossy trees; fronds almost white. 
86. Trichomanes palmatifidum C. Muell. 
Location: C. 
Epiphyte in moss on dwarf trees. Probably more abundant but difficult to 
detect. 
87. Vittaria angustifolia Bl. 
Location: F. 
Epiphyte in mossy forest. 
88. Vittaria elongata Sw. 


Location: L. 
Epiphyte in shady forest. 


89. Vittaria elongata Sw. var. angustifolia Holtt. 
Location: L. 
Epiphyte in forest. 

90. Xiphopteris hieronymusii (C. Chr.) Holtt. 
Locations: A, C, F, H, I, J, K. 
Epiphyte in mossy forest. 

91. Xiphopteris sparsipilosa (Holtt.) Holtt. 


Locations: B, J. 
Epiphyte in mossy forest. 


LisT OF UNIDENTIFIABLE NUMBERS 


92. Asplenium sp., possibly A. pellucidum Lam. 


Location: J. 
Epiphyte in small wet valley near dam. 
Collection. 10.11.74: 1092 (K). 


93. Belvisia sp. 


Location: L. 
Specimen from small valley near dam. 
Collection. 10.11.74: 717] (K). 


94. Ctenopteris sp. 


Location: A. 
Small epiphyte on mossy tree. 
Collection. 12.10.74: 1055 (K). 


42 Gardens’ Bulletin, Singapore — XXX (1977) 


95. Cyathea sp., probably C. obscura (Scort.) Copel. 


Location: J. 
Small sterile plant growing near track. 
Collection. 10.11.74: 7277 (K). 


96. Diplazium sp., near D. pallidum BI. 


Location: J. 
Fertile fronds from side of track. 
Collection. 10.11.74: 7011 (K). 


97. Diplazium sp. 


Location: L. 
Incomplete fertile frond from edge of forest. 
Collection. 19.07.74: 1085 (K). 


98. Hymenophyllum sp., possibly H. polyanthos Sw. 


Location: E. 
Epiphyte on tree in valley. 
Collections. 12.10.74: 1146 & 1147 (K). 


99. Microlepia sp. 


Location: G. 

Single plant with sterile fronds only, much more hairy than M. puberula; 
under observation but lost during road improvements. 

Collections. 9.11.74: 7175 (K) & 1176 (SING). 


100. Trichomanes sp., probably T. maximum BI, 


Locations: H, I. 
Fertile fronds from wet mossy hollows. 
Collections. 2.02.75; 1144 (K) & 1145 (SING). 


101. Trichomanes sp. 


Location: L. 
Sterile frond from mossy forest. 
Collection. 19.07.74; 1248 (K). 


The list of fern species was compiled after numerous expeditions to Gunong 
Ulu Kali over a period of about four years. Records of previous plant collections 
of that place are few. Burkill (1927) indicates that Burn-Murdoch obtained speci- 
mens from the mountain in 1910, and that Robinson sent an expedition to nearby 
Gunong Mengkuang Lebah in 1913. Ridley (1922-25) mentions the conifer 
Dacrydium elatum growing at 2,000 feet on Gunong Ulu Kali, and the Singapore 
Herbarium has a record of one of his collections there in 1914. More recently 
Mrs. Allen (1963) collected ferns at Genting Simpah, a few miles away and at. a 
much lower altitude. And in 1973 and 1975 the staff of the Herbarium, 
Singapore Botanic Gardens, collected some ferns but mainly flowering plants from 
the summit of Gunong Ulu Kali. Further visits could well produce records of more 
and new fern species. 


During recent years development has transformed the mountain peak into a — 
new hill resort. This has resulted in changes in the composition and distribution of 
the flora. In view of the proposed expansion and further development of the Genting 
Highlands it would be interesting to follow these changes due to those in the habitat. 


Ferns of Ulu Kali 43 
Acknowledgements 


I wish to express my gratitude to all who assisted in the preparation of this 
article, and thank: Dr. R. E. Holttum, for examining and critically identifying the 
numerous specimens sent to him, and for his most valuable comments; Mrs, N. P. 
Wong of Genting Highlands Hotel Berhad, for the information she supplied; 
Mr. John Piggott for the photographic records; Encik Othman bin Ali and 
Mr. Philip Khoo, for preparing the art work. 


References 


Allen, B. M. (1963). Ferns of the Quartz Ridges. Mal. Nat. J., 17: 19-32. 


Burgess, P. F. (1969). Ecological factors in hill and mountain forests of the States 
of Malaya. Mal. Nat.. J., 22: 119-128. 


Burkill, I. H. (1927). Botanical Collectors, Collections, and Collecting Places in 
the Malay Peninsular. Gard. Bull. Sing., 4: 113-203. 


Holttum, R. E. (1966). Ferns of Malaya (Revised Flora of Malaya, Vol. IL) 2 ed. 
Govt. Print. Office, Singapore. 


(1971). Studies in the Family Thelypteridaceae III. Blumea, 19. 
17-52. 


Ridley, H. N. (1922-25). Flora of the Malay Peninsula, Vols. I-V. Reeve, London. 


j , ie eae pen 
SP 
ue nm hie , ) 7 Rs 


ae 
é 


wilt to. i 

on} any Jaobi'¢ RU TIA, OTR, QO 

q 4 “AW insets sldeulan 

berlqetpr! sel, erqdienomee lied! abt ner baited | 

bas HA. on 2 i Ao cabo viheqitt 2 
4 i on 1, ait & 


ilu [i ‘she pr 


as unnioF 


My, | ol 
, 5 Wie ad ‘" atv 
y oe A \oM .edebr sta) ort. to vine 


bru Hid ai Holey | tsoigoloott (eet | 
Pe re he 
BO1--Oity Tae Sadie nM 


7 La it 


im ) TES ey " i ‘ tay uf ite w Y teactihoe: a 
Bee Oe tt nt ‘ovtnd TehOes io" 


} ‘all A Ae 
oF ie ' : Aw eS " ' igi 
Gy? iV RPed yA eR eettod ot 


PORnAR thie i 


erorind Rose ee i base on a 
De. 


J tien tek Meise ae) 166 1014, “(ies 


t 


Meek, “ita . 

Kvn fat: ee ae 
Rare Mayeg te Ne Bia Ae 
bei} ers toa Rohunaghe§ Ci 
i PPT Rea f he 
«KI fet oe) ai nang Ay 


~, . ; hed . , hh ‘out ¥ mS mt 
PS, Weare ea ay 
y % ' Se ve ® 
+9 4 y ye I a. 
sin net Py ey ’ a ‘ : . r 
i HRY ay eee a * ie ‘ 


aioe ale eens S69 Byte 
va sete of pve si 


Plate 1. : 
Top left: Acrophorus blumei, part of fertile pinna, x 1. 


; é 7 

Top right: Ctenopteris tenuisecta and orchid, x 4, . ’ 

tiom, left: Cyathea hymenodes, variety with reduced sear ae sa 
Bottom right: Lecanopteris carnosa, apex of frond with soral flaps, 


6 


ate oY. 


Plate 2. 


Top: Ctenopteris contigua, part of fertile lamina, x_ 2. 


Centre: Elaphoglossum sp., fertile (above) and sterile (below) fronds. 
Bottom: Crypsinus laciniatus, fertile fronds, x 4. 


Specific Concept in Humata pectinata (J. E. Smith) Desv. 
by 


G. J. DE JONCHEERE 


Rijksherbarium 
Leiden 


Introduction 


The genus Humata Cav., of Davalliaceous affinity, is a typically tropical East 
Asian group of ferns, extending with a few outlying species to Madagascar in the 
West, Japan in the North and far into the Pacific to the East, but having its centre 
of distribution in Malesia. Thus, several species of Humata occur in Malaya-proper 
and the reader may be referred to the account of the genus in Holttum’s well-known 
book on the ferns of Malaya (1954). 


Holttum remarks on the difficulty of specific delimitation in the genus gene- 
rally, caused by plasticity and variability, also to be observed, by the way, in other 
genera of the same relationship, like Davallia, Scyphularia, etc. In fact, Holttum 
suggests in the elaborate observations he makes under the specific descriptions 
that several species described from adjacent regions are doubtfully distinct from 
the Malayan Humatae recognized by him. But in the case of Humata pectinata 
(J. E. Smith) Desv., additional comment is limited to a short note on its ecology. 


Still, a long history is attached to the name Humata pectinata, also as to how 
it should be interpreted and it may therefore be interesting to follow the vicissitudes 
of the specific concept that have been attached to this name. 


Historical Account 


It was perhaps ironical that J. E. Smith (1793), when first describing Davallia 
pectinata, based his diagnosis on two specimens, collected in localities that could 
hardly have been farther apart. He had received these plants from Banks and after 
the diagnosis Smith added: 


‘Habitat in India Orientali, D, Hurloch 1786, eandem forte in Otaheite 
legit Nelson. H. Banks.”’ 


Alston (1933) commented on these types, still present in the Smith Herbarium 
now at the Linnean Society in London, in a publication that will be more fully 
discussed further on. It should be mentioned at this stage that Alston convincingly 
explained that Hurloch’s plant came most probably from the Nicobar Islands, i.e. 
the most westerly limit of the distributional area of the Humata pectinata alliance, 
whereas the Nelson plant came from the most easterly (Otaheite = Tahiti). 


The lamina of the two types are narrow-deltoid, the lowest segments being the 
largest. However, the basiscopic side of the lowest segments in the Hurloch-collec- 
tion has only one prominent lobe, whereas the Nelson-collection has several, 
gradually passing distally into smaller crenations towards the segmental apex. 

Gaudichaud (1827) described and illustrated Nephrodium gaimardianum, 
typified by a specimen now at Paris from Lawak (= Rawak) Island near Waigeu 
off West Irian, collected by his friend Gaimard. This must be considered an 
entirely independent description: no mention is made of J. E. Smith’s publication 


45 


46 Gardens’ Bulletin, Singapore — XXX (1977) 


and the likeness of N. gaimardianum to Smith’s plant. The fern Gaudichaud 
illustrates is slightly narrower, lanceolate, and has one lobe on the lowest pair of 
segments. 


A year later Wallich published his list (1828) and gave the name Davallia 
parallela to the Singapore collection No. 251. Wallich’s names are “‘nomina nuda’, 
but D. parallela was validated by Hooker, as shown hereunder. 


Wallich’s plant is hardly different from Gaudichaud’s type of Nephrodium 
gaimardianum, with one small lobe on the lowest pair of segments. 


Blume (1828) misconstrued Davallia pectinata J. E. Smith and gave that name 
to a species of Prosaptia, at the same time describing Davallia intermarginalis from 
Java, the type (at Leiden) being closely comparable to Gaudichaud’s and Wallich’s 
types. Blume’s name was recognized as a synonym at an early stage in subsequent 
literature. 


A beautiful plate of Davallia pectinata J. E. Smith was issued by Hooker & 
Greville (1831, pl. 139). The leaf is narrow-deltoid with many basiscopic lobes on 
the lowest pair of segments (“‘pinnae’’). It was drawn from a specimen, collected by 
Menzies in Tahiti and resembles closely J. E. Smith’s Nelson type of the same 
origin. 

It is significant that Hooker at that time had no doubts that Menzies’ plant is 
conspecific with both the syntypes of J. E. Smith, as clearly indicated in the 
elaborate text, accompanying the plate and in which Smith’s type-localities are 
specially mentioned: 


‘‘Hab. In insula Otaheite Menzies. In Malacca et in insulis Nicobar dictis. 
Smith.”’ 


Amongst the above basic descriptions Hooker (1846) made a critical choice in 
his monumental “‘Species Filicum’’. He retained two species, Davallia pectinata 
J. E. Smith and Davallia parallela ‘“‘Wall’’. These taxa were, according to Hooker 
“undoubtedly nearly allied, but distinct’, to be separated by the following 
characters: 


a. D. parallela has the shape of the frond less deltoid, also not so deeply 
divided and therefore never really pinnate, as in D. pectinata. 


b. Margins of segments (pinnae) are horizontally patent and entire, whereas 
in D. pectinata they are spreading and crenate. 


c. Lower margin of lowest pair of segments generally with one solitary lobe, 
rarely more; in D. pectinata pinnatifid with several lobes. 


d. Indusium opening to apex of segments; in D. pectinata opening obliquely 
to the (crenate) margin. 


Apart from these two species, Hooker described Davallia parallela var. f, 
based on Cuming 61 from Luzon (BM, L), previously determined by J. Smith 
(1842) as Humata pectinata (J. E. Smith). It has no lobing of the lowest pair of 
segments, merely auricles. 


As to the nomenclature adopted by Hooker: Wallich’s name was deemed 
acceptable and thus “‘parallela’”’ was given priority over Gaudichaud’s epithet 
“‘gaimardiana”’ which is mentioned in synonymy. Blume’s spurious conception of 
““Davallia pectinata” was not understood (Hooker had not seen Blume’s types). 
“Java, Blume”’ is included in the enumeration of collectors and localities mentioned 
under D. pectinata and at the same time Davallia intermarginalis Blume is mentioned 
as a (doubtfully) separate species, the description from the Enumeratio being merely 
literally cited on the following page of the “‘Species Filicum”’. 


| 


Humata pectinata 47 


That Hooker had his doubts on the origin of Smith’s Hurloch type — and 
by inference on Blume’s “‘Davallia pectinata”” — is indicated by his remark under 
D. pectinata: 


“Sir Jas. Smith gives the East Indies as a locality, on the authority of Mr. 
Hurloch, but perhaps erroneously, for I have never seen it from the Continent 
of India, only from the Pacific’. 


This interesting note also clearly shows that Hooker still considered the 
syntypes of J. E. Smith to be conspecific; moreover, that an initial concept was 
developing in Hooker’s thoughts as to a possible geographical separation of his two 
species. 


At this stage a short review of the generic assignment of the taxa under discus- 
sion would be appropiate. Hooker had a very broad and mostly unnatural conception 
of the genus Davallia, as originally conceived by J. E. Smith. However, in the case 
of D. parallela and D. pectinata, being definitely Davalliaceous ferns, an inclusion 
in Davallia-proper could be defended, especially as Hooker lists these taxa as 
belonging to the subgenus Humata (Cav.). Still, in Hooker & Baur’s (1842) ‘Genera 
Filicum’’, Humata Cav. is recognized as a proper genus, distinct from Davallia 
J. E. Smith, having an indusium that is free at the sides. As this subject is not falling 
within the strict scope of this article, it may suffice to say that in modern literature 
the genus Humata is universally recognized. One must add, however, that it is clearly 
and closely allied to Davallia s.str. and although only differing from Davallia 
virtually in one character, is an easily definable group and perhaps is best regarded 
as a “Genus of convenience’’, to speak with Copeland. 


Continuing the historical review on the subject: in the Synopsis of Hooker 
& Baker (1867) the two species were maintained, but the differentiating characters 
were for the greater part eliminated. In fact, the only clearly definable feature left 
was the lobing of the lowest segments: pinnatifid “with lobes sometimes 4 inch 
long” in Davallia pectinata, whereas in D. parallela “‘the lowest pair (is) sometimes 
auricled”’. Gone are the differences in frond-form (described as ovate-lanceolate in 
both species); the orientation of the sori (oblique in both species); the pinnate or 
pinnatifid condition (being cut down nearly or quite to the rachis in both species). 
Hooker commented under D. pectinata: ‘“‘Quite similar to the preceding 
[D. parallela] in size and texture’. 


D. intermarginalis Blume was no longer mentioned. “‘D. pectinata Blume non 
Smith’ was included in the synonymy of D. contigua var. D. blumei Mett. 
[= Prosaptia] in the Appendix. D. gaimardiana (Gaud.) was mentioned in the 
synonymy of D. parallela as oldest name, but not (surprisingly) adopted. 


The geographical details were given as: D. parallela in Malayan Peninsula and 
Polynesian Islands; D. pectinata in Tropical Polynesian Islands only. 


Hooker’s views had a deciding influence generally in the last century and in this 
particular case maybe up till now. To mention a few examples, Brackenridge (1854) 
recognized Humata pectinata “‘J. Smith” from Tahiti, H. parallela “(Wall)” from 
Samoa, enthusiastically referring to Hooker’s (later discarded) differentiation in the 
“Species Filicum” regarding the orientation of the sori. Presl. (1849) conceived a 
new genus Pachypleuria, differing from Humata Cav. mainly in non-dimorphic 
fronds, and made the new combinations Pachypleura parallela (Hooker) Presl and 
P. pectinata (J. E. Smith) Presl, the former being even assigned by Fée (1852) as 
sole species to yet another genus, Pteroneuron, differing by (pseudo) dorsal sori, 


but neither gave any new ideas on the specific delimitation of the species in 
question. 


The situation seemed to have been consolidated by Christensen (1906) in his 
Index, where he recognized Humata gaimardiana (Gaud.) J. Smith (Syn. Davallia 


48 Gardens’ Bulletin, Singapore — XXX (1977) 


parallela [Wall.] Hooker) and Humata pectinata (J. E. Smith) Desv., the former 
occurring from Burma through Malesia to Polynesia, the latter in Polynesia only. 


There is in this period, however, one notable exception. It was Luerssen (1871) 
who, in his description of the ferns in the Fiji and Samoan Islands, strongly com- 
mented on the weak differentiation given by Hooker between Davallia parallela 
and D. pectinata: 


‘“‘Alle angegebenen, sogar von Hooker und Baker noch festgehaltenen 
Merkmale, welche die Davallia parallela Wall von Davallia pectinata trennen 
sollen, taugen nicht’’. 


which is to say that, according to Luerssen, even the few means of differentiation, 
maintained in Hooker & Baker’s Synopsis, most emphatically do not hold good. 
His elaborate comments clearly point to the many transitions, even on the same 
plant, that exist in the characters, supposed to serve as differential, all being variable 
within the traditional specific delimitation of both taxa. 


Luerssen gave a full and complete list of the synonymy, in which Davallia 
pectinata J. E. Smith is included, but nevertheless adopted the later name Davallia 
gaimardianum (Gaud) ‘‘Presl’’, for which reason is obscure. Presl’s combination is 
even illegitimate according to present rulings, having been published in the Tentamen 
(1836) with a query-mark and as a synonym in the Epimelia (1849). 


Apart from Diels (1899) nobody ever took notice of Luerssen’s view and it 
remained a lone cry in the — taxonomic — wilderness, one might say. 


Christensen’s influence on and significance for modern taxonomic fern-studies 
has been reemphasized quite recently by Holttum (1975). It is therefore no wonder 
that the former’s specific concept on the subject taxa in the Index (1906) and the 
modified version in the Third Supplement (1933) — further discussed below — can 
be retraced in all modern regional floras in the Far East, to mention only: Van 
Alderwerelt’s (1908) Malayan Ferns, Backer & Posthumus’ (1939) Varenflora van 
Java, Tardieu & Christensen’s (1939) Flore Générale de IlIndochine Vol 7, 
Holttum’s (1954) Ferns of Malaya and Copeland’s (1958) Fern Flora of the Philip- 
pines; and as far as the Pacific is concerned: Copeland’s (1929 and 1932) treatises 
on the ferns of Fiji- and Society Islands, Christensen’s (1943) revision of the 
Pteridophyta of Samoa and Brownlie’s (1969) Pteridophyta of New Caledonia. 


In the Third Supplement Christensen also recognized and confirmed several new 
species that had been described in the meantime as belonging to the same alliance. 


Van Alderwerelt van Rosenburgh (1920) proposed Humata lanuginosa from 
Sumatra, syntypes Lérzing 4567, 4764 and Biinnemeyer 3881 (BO, L). In his 
description there is no character that is in any way new to the taxa discussed, apart 
from the profuse scaliness and the presence of hairs on the lower surface of the 
lamina. 


There is another point in Van Alderwerelt’s publication that deserves special 
comment. His types are all from Central Sumatra near Toba Lake, from 
1100-1800 m, from where several other collections have been made, also from Mt. 
Dempo and Sibayak at higher altitudes. This is unusual, as elsewhere, also in 
Sumatra, collections are from the coastal plain or foothills and also often from the 
sea-shore. 


The duplicate of Biinnemeyer’s type-specimen, apparently sent to Leiden some- 
what later, has the epithet “‘lanuginosa’’ deleted and ‘“‘gaimardiana’”’ written instead 
by Van Alderwerelt himself. This tends to show that the author did not believe 
long in his new creation; as several lamina of the syntypes show multiple lobing — 
and going by the criteria that Van Alderwerelt had used in his Malayan Ferns, a 
rectification in Humata pectinata would have been more justified. ; 


; 


Humata pectinata 7 49 


Copeland was more prolific. In his work on the Ferns of the Society Islands 
(1932) he remarks (p. 63): 


‘“‘Apparently in this group of [Humata pectinata] each island has developed 
a peculiar strain which might be construed as a distinct species’. 


but (fortunately) describes and illustrates only two, viz. Humata huahinensis, type 
Grant 5295 from Huahine and H. melanophlebia, type Grant 5144 from Tahaa (not 
seen). This self-imposed restriction is explained on p. 12 where, as on the other 
islands like Moorea and Bora Bora, “‘the characteristic representatives are less fixed 
in their pecularities and are accordingly left without distinctive names’’. 


Humata huahinensis is described by Copeland as being very near to H. pecti- 
nata, but having a black stipe and deviating towards H. gaimardiana by being 
pinnatifid and position of sori, also the closely placed segments. 


As to Humata melanophlebia, Copeland comments that it is like H. huahinensis, 
but pinnate with more remote pinnae (or segments more distally) which are clearly 
lobed, mainly basiscopically, the segments becoming inciso-serrate. Copeland (on 
page 63) mentions Grant’s collection 5144 [= type of H. melanophlebia] again 
under H. huahinensis. Be that as it may, the proposed elevation to “‘species” of 
these Pacific representatives does not thereby become more convincing. 


In 1940 Copeland described and illustrated another species, Humata tenuivenia, 
type Brass 14082 from New Guinea: 


“‘Humata pectinatae affinis, venis tenuibus inconspicuis, soris perlatis curvis, 
indusiis brevibus destincta’’. 


The (iso)types (L and BO) and the photograph show these specimens to be 
robust plants from a shady wet habitat (“low epiphyte in coastal swamp-forest’’). 
Copeland specially comments on the curved sori which can face the margin and 
apex or, in extreme cases, is curved so far that it faces at an angle to the costa too. 


A few years earlier Copeland had been the instigator of yet another name. He 
had requested Alston to have a new look at J. E. Smith’s types of Davallia pecti- 
nata. As a result, Alston (1933, l.c.) lectotypified Davallia pectinata J. E. Smith on 
the Hurloch specimen, dispersing Hooker’s previous doubts as to the origin by 
showing that it came most probably from the Nicobar Islands and in any case from 
Western regions, not from the Pacific. Arguing further, Alston came to the con- 
clusion that the name Humata gaimardiana (Gaud.) J. Smith (of Christensen’s 
Index with a predominantly Western distribution) must in fact be replaced by 
Humata pectinata (J. E. Smith) Desv. and that the other syntype of J. E. Smith 
(Hooker’s and Christensen’s Humata pectinata with a Pacific distribution) had no 
name. Alston thus reversed the ideas hitherto current on geographical distribution, 
but maintained the traditional separation of two specific entities. 


The new name Alston introduced was Humata banksii and the typification is 
based on the Tahiti collection No. 1769 (BM) by Banks, ‘‘Nelson’s specimens being 
poor, both in the herbaria Smith and Banks’’. Alston’s description emphasizes 
“profundae pinnatis’”’ and “‘inferioribus margine inferiore pinnatifidis’’, otherwise 
does not give any further characters of differentiation from his H. pectinata. 
Actually Alston followed the description as included by Hooker & Baker in the 
Synopsis for (their) Davallia pectinata. Completely absent are comments on notably 
geographical distribution, or a comparison with e.g. Copeland’s previously described 
species (1932) of which H. Auahinensis is practically identical with Banks’ plant. 
It is, as if Alston was wary to incriminate himself further in sorting out the 
antecedents of his new proposal. 


50 Gardens’ Bulletin, Singapore — XXX (1977) 


Nevertheless, Alston’s views, whether stated or only implied, were immediately 
followed by Christensen (1933) in the third Supplement of the Index and con- 
sequently in all the more local Flora’s mentioned previously: Western plants now 
indiscriminately being called Humata pectinata (J. E. Smith) Desv., Eastern (Poly- 
nesian) plants referred to Humata banksii Alston. 


But, Copeland did not agree. His comment on the Humatae in the “‘Oleandrid 


Ferns of New Guinea’”’ (1940) may be appropriately quoted in full at the end of this 
historical survey: 


“This is the largest and most difficult genus of the group. Typically 
epiphytes, the individuals are subject to wide variations in exposure, and 
some of them are very responsive to these differences. Independent of the 
environment, some species seem to be notably variable ... Smith combined 
a Malayan and a Tahitian plant in describing Davallia (Humata) pectinata. 
H. parallela (Wall) Brack. and H. gaimardiana (Gaud.) J. Sm. were sub- 
sequently described in the same group. With many specimens from the 
Society Islands, it seemed to me that each island had its own, more or less 
distinct form, none of these like the comparatively uniform Malayan plant. 
It occurred to me that the Malayan plant might be the real H. pectinata, 
so I invited comparison of types by Mr. Alston. The result was his finding 
the Tahiti specimens [sic!] to be of two species, H. pectinata and H. banksii. 
It is my conclusion, not his that the Malayan plant must be H. parallela, for 
I have no Malayan specimen duplicated by any from Tahiti. 


Some characters which usually serve as specifically diagnostic, serve so badly 
in Humata. Size varies greatly as a matter of plasticity (response to environ- 
ment). So with size does the dissection of the frond; and so probably do 
texture and laxness. Paleae are likely to be deciduous. And dimorphism is 
subject to some reversion. 


In the light of the foregoing discussion it will be understood that considerable 
work on this genus leaves me ill-satisfied. The presentation here given is the 
best I can make with the present material’. 


One could have full sympathy with this lament, were it not that in the ensuing 
key, size, dissection of frond, degree of scaliness and dimorphism figure promi- 
nently as differentiating characters. Of the 21 species, Copeland distinguishes in 
New Guinea alone, not less than six are described as new, of which one, Humata 
tenuivenia falls within the affinity of the taxa under review, as discussed above. 


Copeland’s confused rejection of Alston’s interpretation of J. E. Smith’s types 
was, for that matter, corrected later (1958) in the Fern Flora of the Philippines, 
where he uses the name Humata pectinata (J. E. Smith) Desv. for Philippine 
specimens, mentioning Davallia parallela ‘“‘Wall’”’ as a synonym. 


Observations 


When looking back on the features that have been used to distinguish between 
the taxa, as discussed heretofore, a hard core of only two characteristics remain, 


viz. the lobing of lowest pair of segments and the pinnatifid against the pinnate 
condition. 


As to the last feature, the wing alongside the rachis is always distally broader 
than below and is, even in the most deeply incised leaves, never entirely absent. 
One cannot talk therefore of a real pinnate condition, it is always pseudo-pinnate 
and this variable point of distinction is in reality non-existent. 


Humata pectinata 51 


As to the (basiscopic) lobing of the lowest pair(s) of segments, transitions can 
be found on the same rhizome sometimes, from non- one-, two- to multi-lobed. A 
clinal pattern can be observed, whereby the multilobed basal segments become less 
frequent from East to West (nearly absent in Malaya and the Philippines), whereas 
the non-lobed or auricled/one lobed segments decrease from West to East, to being 
virtually absent in the Eastern Pacific. 


The same pattern of gradual transitions can be found in the length of the lowest 
segments, being either longer, equal to, or shorter than the next pair, this making 
the overall leaf-shape either narrow deltoid, lanceolate/linear or narrow-ovoid. The 
sinuses vary in width from nearly equal to the width of the segments to mere slits, 
variable even on the same leaf or plant. 


The above observations make differences in leaf-morphology unfit for the 
proper distinction of species and the recognition of Humata banksii Alston (and 
Humata huahinensis Copel.) seems to be based on weak ground. 


Humata melanophlebia Copel. is an extreme case, where the lobing is also 
conspicuous on the next lower pair(s) of segments and more or less extended to the 
acroscopic side of the segments as well. 


Differences in scaliness have hardly been used in proposing separate taxa, the 
variability in caducity and density having been generally understood. Van Alder- 
werelt’s “‘hairs’’, described in Humata lanuginosa, are nothing but the highly 
dissected, hyaline squamules generally found — but not so obvious — on the 
underside of the lamina. 


Actually, often the scales are not, or not well described, a glabrous or sub- 
glabrous condition prevailing in mature leaves. But young leaves (and stipes) are 
always scaly and the reader may be referred to Appendix I, where a full description 
of the scales can be found. 


The venation is always + flattened, conspicuous below, + parallel with none 
or 1-2 forkings, 3 forkings only occurring in very large leaves, as in Humata 
tenuivenia Copel. The ancillary veinlet round the (terminal) sorus, basis for Fée’s 
genus Pteroneuron can also be absent, or the veinlet is forked far below the sorus, 
thus becoming just a part of the normally forked venation, Transitions can be 


found even on the same segments. 


The position and orientation of the sorus is also inconstant. It is always 
intramarginal and the indusium can be semi-circular to crescent-shaped, in extreme 
cases becoming reniform (Humata tenuivenia Copel.). Distally the sorus mostly 
faces the apex of the segment, lower down (obliquely) the margin. 


Conclusion 


The above review was made in the framework of a general study of the genus 
Humata Cay. — and other davallioid genera — the results of which will eventually 
be published in the Flora Malesiana. As such, Van Steenis’ forceful, but lucid 
essay on “Specific Delimitation” is here considered an excellent guide to extricate 
oneself from further confusion. In itself the present study is an illustration of 
practically all the pitfalls, so convincingly summed up by Van Steenis, that can 
beset taxonomy and although his work is mostly based on vast experience in 
phanerogamic taxonomy, it is certainly also fully applicable to ferns. 

Therefore, the author cannot but come to one conclusion, viz. that all the taxa 


under discussion can best be regarded as one polymorphic species with a wide 
distribution, for which the name Humata pectinata (J. E. Smith) Desv. should be 


52 Gardens’ Bulletin, Singapore — XXX (1977) 


used. Luerssen’s views of nearly a hundred years ago are thus herewith fully 
confirmed. 


Delimitation from allied species like Humata repens (Linn. f.) Diels, H. vestita 
(BI.) Moore and H. heterophylla (J. E. Smith) Desv. gives no difficulty and shall 
not be further elaborated on in this context. 


Recognition of infra-specific taxa is omitted; the variable, transitional (and 
clinal) nature of the characteristics, mentioned hitherto make them unsuitable for 
the delimitation of recognisable subspecies or varieties. 


The geographic distribution of Humata pectinata as here construed is a clear- 
cut case of dependency on certain climatological circumstances. The area covers 
practically exactly and totally the zone of small seasonal variations in temperature 
(up to 35° F = 20° C) in the Eastern Tropics and Pacific, with the exception of 
that part of the zone which is liable to an appreciable monsoon. This explains the 
striking absence of the species from (East) Java, South-Sulawesi, Lesser Sunda Isles 
and Southern Moluccas in an otherwise continuous distribution throughout Malesia 
and Polynesia up to Tahiti. New Caledonia in the South and Taiwan in the North, 
in both of which islands the species is rare, are only just outside the zone as above 
referred to. Climatological maps covering the region can be found in Goode (1947). 


A special note should be devoted to the occurrence of the species in Java. 
Although Blume’s type of Davallia intermarginalis came ostensibly from Western 
Java, no further collections — apart from an old Junghuhn specimen at BO — have 
been made in that area for nearly 150 years. Nevertheless, potentially West Java 
(as against the eastern part of that island) could harbour our species, having only 
a weak monsoon. Could it have become extinct there by the vast destruction of 
lowland forest of that island? 


The high-altitude collections in Central Sumatra have already been commented 
upon. This could be a special ecotype, but from New Guinea specimens from 
medium elevations (1100 m) are also known. 


It could be argued that research of a karyological nature, especially cyto- 
genetical, could produce better substantiated and more refined results than is 
possible by limiting the study to only herbarium material. Whilst this limitation 
is recognised it is regretted that such an approach is precluded by the unavailablity 
of live material. Whether it would really help in tackling the problem remains to 
be demonstrated. The mere thought of the difficulties involved in obtaining live 
plants from the whole distributional area and the time-consuming organization in 
rigging up a hybridization programme suggests that the above reserve if not illusory 
would, in practice be hard to tackle in a satisfactory way. 


Finally this relatively uncomplicated, but nevertheless perforce lengthy account 
on how the species Humata pectinata has to be construed can be equally repeated 
for other taxa of the genus. As the publication of such detailed discussions would 
lead too far and be a tedious repetition, this study may serve as an example as 
to how the specific concept in the forthcoming treaties of the whole group will be 
maintained. That a considerable reduction in names will result — as already 
indicated by Holttum (1954) — is certain. 


Acknowledgments 


To the keepers of the Herbaria at Bogor, British Museum and Kew, I would 
like to express my gratitude for cooperation and assistance extended. To my friend 
E. Hennipman of Leiden Herbarium I am thankful for reading the script and his 
sound advice. 


Humata pectinata 


Bibliographical References 


v. Alderwerelt v. Rosenburgh, C. R. W. K. (1909) Malayan Ferns: 284-291. 
. (1920) Bull. Jard. Bot. Bzg. 3/2: 155. 

Alston, A. H. G. (1933) Phil. Journ. Sc. 50/2: 175-176, t. 1 fig. 1. 
Backer, C. A. and Posthumus, 0. (1939) Varenflora Java: 96. 

Blume, C. L. (1828) En. Pl. Javae, Fil.: 229, 230. 

Brackenridge, W. D. in Wilkes, C. (1854) U.S. Expl. Exp. 16: 229. 
Brownlie, G. (1969) Fl. Nouv. Cal. 3: 149. 

Christensen, C. (1906) Index Fil.: 353, 354. 

. (1934) Suppl. 3 (1917-1933): 112. 

. (1943) Pter. Samoa, Bish. Mus. Bull. 177: 37. 

Copeland, E. B. (1929) Ferns Fiji, Bish. Mus. Bull. 59: 89. 

. (1932) Pter. Soc. Isl., Bish. Mus. Bull. 93: 11-12, 62-63, t. 12. 
—— . (1940) Phil. Journ. Sc. 73/3: 350, t. 3. 

. (1953) Fern. Fl. Phil.: 178. 

Diels, L. (1899) Nat. Pfl. 1/4: 209. 

Fée, A. L. A. (1852) Gen. Fil.: 320, 321, t. 25B fig. 1. 
Gaudichaud-Beaupré, C. (1827) Freyc. Voy. Bot.: 335, t. 12 fig. 1. 
Good, R. (1947), Geography FI. Plants: t. 21, t. 24. 

Holttum, R. E. (1954) Ferns. Mal.: 364~371. 

. (1975) Taxon 24/4: 499-500. 

Hooker, W. J. (1846) Spec. Fil.: 153, 154, t. 42A. 

Hooker, W. J. & Baker, J. G. (1868) Syn. Fil.: 89. 

Hooker, W. J. & Bauer, F. (1842) Gen. Fil.: t. 114A. 

Hooker, W. J. & Greville, R. K. (1831) Ic. Fil.: t. 139. 

Luerssen, C. (1871) Fil. Graeff., Mitt. Gesamtg. Bot. 1: 206-207. 

Presl, C. B. (1836) Tent. Pt.: 79, 128. 

. (1849) Epim. Bot.: 98. 

Smith, J. (1841) Journ. Bot. 3: 416. 

. (1842) London Journ. Bot. 1: 425. 

Smith, J. E. (1793) Mém. Ac. Tprin 5: 415. 

v. Steenis, C. G. G. J. (1957) Fi. Males. ser. 1, 5/3: CLXVII-CCXXXIV. 
Tardieu-Blot, M. L. & Christensen, C. (1939) Fl. Gén. Indo-Chine 7/2: 109. 
Wallich, N. (1828) List: No. 251. 


54 Gardens’ Bulletin, Singapore — XXX (1977) 


APPENDIX I 
NOMENCLATURE 


Humata pectinata (J. E. Smith) Desv. (1827). Prodr.: 323. — Davallia pectinata J. E. 
Smith (1793) Mém. Ac. Turin 5: 415; Trichomanes Poir. in Lam (1808) Encycl. 8: 78; 
Pachypleuria Presl. (1849) Epim.: 98 — Type: Hurloch/Soc. Unit. Frat. s.n. from Ind. Or. 
(probably Nicobar Isl.) as lectotypified by Alston (1933) Phil. Journ. Sc. 50: 175, t. 1 fig. 1. 


Nephrodium gaimardianum Gaud. (1827) Freyc. Voy. Bat.: 335, t. 12 fig. 1; Nephrolepis 
Pres]. (1836) Tent.: 79; Humata J. Smith (1842) London Journ. Bot. 1: 425; Davallia [Presl. 
(1836) Tent.: 128 (comb. ill.)] Kuhn (1869) Zool. Bot. Ges. Wien: 580 — Type: Gaudichaud 
(Gaimard) s.n. from P. Lawak = Rawak/Waigeu/W. Irian. 


Davallia parallela [Wallich (1828) List No. 251 (nomen nudum)] Hooker (1846) Sp. Fil.: 
153, t. 4248; Pachypleuria Presl. (1849) Epim.: 98; Pteroneuron Fée (1852) Gen. Fil.: 320, 
t. 256; Humata Brack. in Wilkes (1854) U.S. Expl. Exp. 16 Bot.: 229; Oleandra Keys. (1873) 
Pol. Cyath. Herb. Bung.: 41 — Type: Wallich 251 Singapore. 


Davallia parallela var. 8 Hook. (1846) Sp. Fil.: 153 — Type: Cuming 61 Luzon. 


Davallia intermarginalis Blume (1828) Enum.; 230; Pachypleuria Presl. (1849) Epim.: 98; 
Humata Moore (1861) Index: 296 — Type: Blume s.n. Java. 


Humata lanuginosa v.A.v.R. (1920) Bull. Jard. Bot. Bzg. 3/2: 155 — Syntypes; Lérzing 
4567, 4764, B. Baroe/Sumatra; Biinnemeyer 3881 G. Malintang/Sumatra. 


Humata huahinensis Copel. (1932) Bish. Mus. Bull. 93: 11. t. 126 — Type: Grant 5295 
Huahine/Soc. Isl. 


Humata melanophlebia Copel. (1932) Bish. Mus. Bull. 93: 11, t. 128 — Type: Grant 
5144 Tahaa/Soc. Isl. 


Humata banksii Alston (1933) Phil. Journ. Sc. 50: 176 — Type: Banks 1769 Tahiti/Soc. 
Isl. 


Humata tenuivenia Copel. (1940) Phil. Journ. Sc. 73: 350, t. 3 — Type: Brass 14082 
Idenburg R./W. Irian. 


DESCRIPTION 


Rhizome upto several metres long-creeping, slender (14-24 mm), blackish with white- 
chalky patches, dictyostelic and dorsiventral, vascular tissue much dissected with two main 
bundles; bearing articulated fronds, on short phyllopodia, 2-5 cm or more apart; thickly 
set with appressed. imbricated scales, 4 to 5 mm long, + 1 mm wide at the oblong peltate 
base, from there tapering to the acute apex, castaneous, the thin margin often hyaline /whitish, 
when young bearing marginal crinkly hairs which are mostly soon deciduous, the edge becom- 
ing entire at an early stage. 


Stipe thin (1 mm g) but firm, as long as, or somewhat shorter than the lamina, green 
to reddish brown when living, brown to blackish when dried, sulcate; vascular tissue with 
mostly three bundless, becoming fused upwards; loosely set with caducous scales and often 
entirely glabrous with age; scales like those of rhizome but not so appressed and more 
rounded, becoming smaller, paler, the edge more dissected. 


Lamina firm, coriaceous, (5) 7-18 cm long, 4-6 (8) cm wide, narrowly deltoid, lanceolate / 
linear to narrowly ovate, dark green above, paler below, pinnatifid/pectinate with truncate/ 
cordate base and short coadunate apex; segments many (12-30) which are sessile and 
confluent by a narrow wing along the rachis; wing occasionally inconspicuous between the 
lowest pairs of segments (the lamina then becoming pseudo-pinnate) but always widening 
upwards; sinuses deep and narrow, to about as wide as the pinnae but usually much 
narrower, becoming distally shallower towards the crenate to entire apex; rachis prominent, 
green to brown/blackish, sulcate except at the apex; scales loosely covering the young 
unfolding leaves, equal to those on the stipe, however, especially on the lamina becoming 
even more hyaline and dissolute, variously deciduous but mostly leaving a few traces on 
+ See surface of rachis and costa, as well as on the margins of the often (sub) glabrous 
amina. 


Segments to 6 (8) mm wide, straight and horizontally patent or a little curved upwards, 
the apex bluntly acute to rounded; edge in barren leaves rarely quite entire, commonly 
shallowly crenate/sinuate at apex only, less often the incisions becoming more pronounced 
and continuing to bottom of sinus; in the very slightly and inconspicuously contracted fertile 
leaves incisions more prominent; the lowest pair of segments variable, either somewhat 
shorter or equal (ovate-lanceolate form), or if more developed, becoming longer than the next 
pair of segments (narrow-deltoid form), the basiscopic edge being either entire, auricled, 
often prominently, or two/multi lobed, the lobes grading to the crenated apex, the lowest 
segments then becoming unilaterally pinnatifid; venation, apart from the prominent costa, 


Humata pectinata 55 


distinct on lower surface only, the veins coarse, brown and parallel, not or 1, 2, (3) times 
forked, both upper and lower basal primary veins springing from very base of costa or in 
larger fronds the lower directly from the rachis. 


Sori terminal on the swollen acroscopic (when forked) vein-ending, forming an intra- 
marginal, often crowded row; away from the apex often with an accessory posterior veinlet 
running round the sorus to just within the leaf-edge; indusium brown, + | mm wide, finely 
striated, firm and permanent, semi-circular or crescent-shaped to reniform, the lower straight 
to convex side attached in the middle for + half its length, otherwise the indusium free, 


opening to the apex or (especially lower down) obliquely to the leaf-edge; sporangia with 
12-14 indurated cells; spores monolite, verrucate-rugulate, 20 x 13 4 with crenulated margin. 


GEOGRAPHICAL DISTRIBUTION 


From Southern Burma (Mergui), Lower Thailand, South Viet Nam, and Taiwan, 
throughout Malesia with the exception of those regions with an appreciable monsoon (Middle 
and Eastern Java, Lesser Sunda Islands, South-Sulawesi and South Moluccas), extending 
further into Tropical Polynesia to Society Islands in the East and New Caledonia in the South. 


ECOLOGY 


Epiphyte, high and low, on trees; or terrestrial on boles and rocks, in often exposed 
habitats near seashore and in light forest, steep banks, even padangs on bare sand, but also 
known from swamp-forest in shade. Can apparently stand a lot of exposure, the leaves 
curling up in dry periods [Holttum (1954)]. From sea-level to 800 m, going up to 2000 m 
in Sumatra and to medium altitudes in New Guinea. 


APPENDIX II 
ILLUSTRATIONS 


Various facies of Humata pectinata (J. E. Smith) Desv. 
1. Sinclair, S. F. 40582 (1955) North side of Cape Rochado-Malacca- Malaya. 


“Rocky wooded seashore. Creeping rhizome on sea cliffs. Fruiting. Fronds curled 
up with the drought. Flattened by immersing in water”. 


Two leaves from same rhizome. 


2. Lérzing 15533 (1929), East Mt. Sibayak-Sumatra-Indonesia, 1300-1440 m. 


“Primary forest, epiphytic, and, in lighter places, on prostrate trunks and rocks. 
Not rare”. 


Two fronds from same rhizome. 


3. Liitjeharms 4674 (1936), Enggano off West Sumatra-Indonesia. 
“Epiphytisch op klapperstammen. Strand bij Kiojoh”. 
Small leaves on rhizome. 
4. Van Niel 3383 (1964), Tutong-Brunei-North Borneo. 
“Under clumps of trees in an open vegetation (sand)”. 
Two leaves on same (branched) rhizome. 


5. Cuming 61 (1836), Luzon-Philippines. 
No further details — Type Davallia parallela var. § Hooker. 
One leaf from isotype at L. 


6. Brass 14082 (1939), Bernhard Camp-Idenburg River-W. Irian-Indonesia. 


“Low epiphyte in flooded rain forest of rive-plain at 50 m.” — Type Humata 
tenuivenia Copel. 


One leaf from isotype at L. 
7. Van Balgooy 1983 (1971), W. slope Pahia-Bora2-Society Islands, 300 m. 
“Epiphyte, rootstock creeping”. 
One large, one small leaf from same collection. 


56 


Gardens’ Bulletin, Singapore — XXX (1977) 


37 


Humata pectinata 


58 


Gardens’ Bulletin, Singapore — XXX (1977) 


Curcuma zedoaria 


B. L. BuRTT 


Royal Botanic Garden, Edinburgh 


My previous notes on the nomenclature of Curcuma zedoaria (Notes Roy. 
Bot. Gard. Edinburgh, 31: 226. 1972) were inaccurate. At that time I pointed out, 
correctly, that ‘“‘Amiomum zedoaria Berg.”’, which is always cited as the basionym 
of Curcuma zedoaria, has no existence as a binomial, I then said that the first 
valid publication of the name Amomum zedoaria was by Plenck in 1789, by which 
time the plant had already been named A. Jatifolium by Lamarck (1783). Where 
I, and others, have erred till now is in failing to notice that Amomum zedoaria was 
validly published in the German version of Houttuyn’s Natuurlijke Historie by 
Christmann & Panzer (1779). Details of this work and its relation to Houttuyn’s 
original publication are given by Merrill (in Journ. Arn. Arb. 19: 291. 1938). 
Briefly, the important point for our present purpose is that the work by Christmann 
& Panzer is not a literal German translation from Houttuyn: it incorporates both 
changes and new matter. Merrill listed all the new names he observed, but 
unfortunately Amomum zedoaria in the German version escaped him, and, not 
having the original work at hand, this helped to lull me into a sense of security. 
Although the work as a whole may be attributed to Christmann & Panzer, Merrill 
showed that new names appearing in volumes 1-7 should be attributed solely to 
Christmann. 


Christmann cites a number of authors under Amomum zedoaria: Bergius, 
Rheede, Rumphius, Petiver and others. It is the description by Rheede under the 
name Kua which gives the most detailed and reliable account of this plant: this 
was cited by Bergius at the head of his references and formed the basis of 
Lamarck’s Amomum latifolium and it is Rheede’s illustration of it that was re- 
drawn for Plenck’s plate. There is every reason for taking this reference as the 
“lectotype” of the name Amomum zedoaria Christm., and this I accordingly 
propose. — 

Amomum zedoaria was published by Christmann in 1779 and it thus provides 
the earliest valid post-Linnaean epithet for a Curcuma. Its transfer to that genus 
may still be attributed to Roscoe, for Roscoe cites Willdenow and Willdenow gives 
the reference “‘Houtt. Linn. Syst. 5, p. 12” which proves to refer to the German 
edition of Christmann & Panzer. It may be noted that in Houttuyn’s original 
Dutch edition (Handl. Pl. Kruidk. 7: 10. 1777) zedoary is referred to Amomum 
zerumbeth, which is Zingiber zerumbet (L.) Roscoe. 


As the basis for the name Curcuma zedoaria is Rheede’s description and plate 
it has seemed not without interest to supply a translation and reproduction. 


Kua 
Plate VII 


Kua, in Brahmi Acua, is the special and common name for all those plants 
that ought to be referred to species of ginger; they are eight in number. Kua, 
Tsjana Kua, Mallan Kua, Manga Kua, Mangella Kua, Intsyi Kua, Katou-Intsyi 
Kua. The first species likes all soils, especially the sandy ones in which it grows 


59 


60 Gardens’ Bulletin, Singapore — XXX (1977) 


spontaneously and spreads. The rhizome runs cut horizontally from a single centre, 
white, thick, tuberous, with slender fibrous rootlets here and there, firm, run 
through with many whitish threads [i.e. vascular bundles], with glabrous skin; it 
is divided along its length into tuberous segments about two fingers thick and a 
span [c. 19 cm] long, branched, and filled with a clear juice. The taste is sharp, 
pinching the tongue, irritant, aromatic, The scent is strong and pleasant as if many 
aromas were mixed together. The leaves arise from the ends of the rhizome, up 
to a cubit [c. 46 cm] long and two spans [c. 38 cm] wide in the middle; acuminate 
at the tip, narrowed at the base, with a prominent midrib; this sends out many 
lateral veins, running transversely forward, equally spaced; they disappear at the 
extreme edge where they embrace the margin of the leaf with finer veinlets; the 
inner part of the leaf is somewhat folded and reflexed, marked with grooves. The 
colour on the inner part is dark, on the outer paler. For the rest the leaves are 
glabrous, shining, clear. The stems are a foot and a half high, full of greenish 
pith, but no shoots are to be seen on them; however, from the rhizome a special 
flowering stem arises, covered by a number of long leafy scales very closely 
enfolding it; and these leaves are more than a digit long, broadened upwards and 
then again contracted to a small point, with the tips bent back from the scape; 
they are glabrous with slender longitudinal veinlets which, unless they be broken, 
are nearly invisible; the Jower ones are green, the middle ones yellow, the upper 
red purple or light blue, the uppermost becoming almost white, pretty, sweet-smell- 
ing, and tasting like the root. The flowers are produced in the gaps between the 
scales [i.e. bracts], 2 or 3 at a time, bell-shaped, the lower part and inside whitish, 
thin, enclosed in a transparent membrane [i.e. calyx], 6-petalled [i.e. with 3 petals 
and 3 petaloid staminodes] the upper one pellucid, shaped like a helmet, erect; 
they are yellow ana white, folded, and scarcely to be seen unless the whole flower 
is separated from the stem and in turn taken apart itself, when one is seen to be 
bent down to a fimbriate margin; in the middle the flower produces a thread 
[i.e. the style] provided with a spur like a dragon’s head, and recalling a little 
winged bird with erect tail [this refers to the spurred anther between the lobes of 
which the upper part of the style is held]. The scent is pleasant, but when the 
flowers are broken from the pouch and bruised between the fingers they have a 
similar smell to the leaves, and a rather bitter flavour. The seed capsules are small 
and round, depressed, and contain grey seeds like those of Tsjana Kua, but they are 
rarely found. The rhizome is long persistent; the leaves die down and shoot forth 
again in July and August. The grated roots, washed several times, leave a flour 
much valued by the indigenous peoples who make a porrage from it. A potion of 
bruised root and fresh juice arrests all inflammation of the intestines, purges the 
kidneys, stops white flux, cures gonorrhea and purges the blood. The juice of the 
leaves is drunk by people with dropsy, it takes away the swelling of the abdomen, 
is a moderate laxative, and expels the viscid slime of the intestines. 


[Footnote by Commelin.] Kua, according to the opinion of the illustrious Paul 
Hermann is the garden Zerumbeth of Garcia [da Orta], whose description like that 
of the Arabic authors is rather obscure; by the Singhalese it is called Walinghura, 
that is wild ginger, the Malabars call it Kua. The opinion of Hermann agrees very 
closely with that of our author. 


The other plants mentioned by their local names, of which the spelling varies 
slightly in different parts of the work, are: — 


Tsjana Kua — Costus speciosus (Koenig) Sm. 
Mallan Kua — Kaempferia rotunda L. 

Manga Kua — Boesenbergia rotunda (L.) Mansf. 
Mangella Kua — Curcuma longa L. 

Intsyi Kua — Zingiber officinale (L.) Rose. 
Katou-Intsyi Kua — Zingiber zerumbet (L.) Rose. 
Mala-Intsyi Kua — Alpinia allughas (Retz.) Rosc. 


4 uy . § 

A - a e bs at 

=a i ie 
eek A eee ie 


oh ES? Rerniny Ae 
- < vies 1 ee gh wet 


at eae ae 


; ee 
<2 


Curcuma zedoaria | 61 


Rheede’s description does not, of course, give the precise technical detail that 
we look for to-day; for that we turn to Holttum’s account of the Zingiberaceae 
of the Malay Peninsula (this Bulletin, 13: 71. 1950). Rheede, however, supplies a 
remarkably vivid account of a plant whose whole structure was a novelty at that 
time. It has not seemed justifiable to reprint the original latin, but those interested 
in the development of botanical terminology may be interested to note that in 
it calyx refers to the cup or pouch formed by the bract, filamentum to the thread- 
like style, and pollen to the flour obtained from the root. 


I have referred throughout to Rheede. Hendryk van Rheede tot Draakestein 
was appointed Governor of the Dutch possession in Malabar in 1667. He organized 
the bringing together of the materials that were sent back to Holland and published 
as the Hortus Malabaricus. Burkill (Chap. Hist. Bot. India, pp. 6-7, 1965) tells 
us that the drawings were made by an artist-missionary Matthaeus, that the accounts 
of the plants were rendered into Portuguese by an interpreter and thence into latin 
by the secretary of the local government Hermann van Doner. Then drawings and 
descriptions were assembled by Johannes Casearius and were sent to Holland for 
publication, where they were edited partly by A. Seyn but largely, after Seyn’s 
death, by J. Commelin who was in charge of the Hortus Medicus at Amsterdam. 


Curcuma zedoaria is a species that has been in cultivation for a very long 
time and its natural origin has never been precisely established. This is a parti- 
cularly difficult problem as it very easily becomes naturalized. In fact it is on 
record that although C. zedoaria very rarely flowers under conditions of cultivation, 
it does so freely where it runs wild. It is now found in most parts of India and 
south-east Asia. Like other members of the genus it dies down completely after 
flowering and has a resting phase: which at least suggests that its origin was in 
the monsoon areas rather than in Malaya. If, as often stated, it originated in N. E. 
India, this would be a good focal point for its spread south into peninsular India, 
east to China and SE to Malaya. 


Synonymy and useful references are detailed below; those appearing in square 
brackets are not nomenclaturally valid names. 


Curcuma zedoaria (Christm.) Roscoe in Trans. Linn. Soc. 8: 354 (1807) et 
Monandr. Pl. Scitam. t. 109 (1825); Horan., Monogr, Scit. 23 (1862); Baker in 
Hook, f., Fl. Brit. Ind. 6: 210 (1890); Trimen, Handb. Fl. Ceylon, 4: 241 (1898); 
K. Schum., Pflanzenr. Zingiber. 110 (1904); Gagnepain in Lecomte, Fl. Gen. Ind. 
Chin. 6: 67 (1908); Merrill, Enum. Phil. Fl. Pl. 1: 243 (1924) et in Trans. Amer. 
Phil. Soc. N.S. 24 pt. 2: 119 (1938); Burkill, Dict. Econ. Prod. Mal. Pen. 714 
(1935); Holttum in Gard. Bull. Singapore 13: 71, fig. 5 (1950); Wealth of India 
2: 405 (1950); Backer & Bakh. f., Fl. Java, 3: 71, 72 (1968). 


Syn: [Kua Rheede, Hort. Malab. 11: 13, t. 7 (1692)] 


[Zedoaria Camellus, Herb. Stirp. Luzon, Syll. 23 No. 9 in Ray, Hist. Pl. 3 
App. (1704).] 

[Zedoaria officinarum Petiver, Gagophyl. Nat. dec. 3, 5, tab. 23 f. 1 
(1704-67).] 

[Zerumbed vel Tommon [primum] Rumph., Herb. Amboin. 5: 169 excl. t. 
68 (1747) — fide Valeton in Merrill, Interp. Herb. Amboin, 164 (1917).] 
[Amomum scapo nudo, spica laxa truncata Bergius, Mat. Med. 4 (1778), 
ed. 2, 4 (1782).] 


Amomum zedoaria Christm. in Christm, & Panzer, Linn. Pflanzensyst. 
5: 12 (1779); Plenck, Ic. Pl. Med. 2: 12, t. 11 (1789); Willd., Sp. PI. 
1: 7 (1797). 


Amomum latifolium Lam., Encycl. 1: 134 (1783). Type: Kua Rheede. 


62 


Gardens’ Bulletin, Singapore — XXX (1977) 


Curcuma pallida Lour., Fl. Coch. 9 (1790) et ed. Willd. 12 (1793). Type: 
‘“‘in agrestis in Cochinchina et Cantone Sinarum’”’ (no specimen known). 

Curcuma zerumbet Roxb, in Asiat. Research, 11: 332 (1810) et Pl. Coro- 
mand. 3, t. 201 (1819); Alston in Trimen, Handb. Fl. Ceylon Suppl. 281 


(1931); Burtt & Smith in Notes R.B.G. Edin, 31: 203 (1972) sub Curcuma 
(Erndlia subpersonata Giseke); nom. illegit. 


Curcuma speciosa Link, Enum. Pl. Hort. Berol. 1: 5 (1821); nom. illegit. 


Studies in the Systematics of Filmy Ferns 
II. A note on Meringium and the taxa allied to this 


by 
K. IWATSUKI 


Department of Botany, Faculty of Science 
Kyoto University, Kyoto, 606 


Japan 


Summary 


The evaluated taxonomic features of Meringium and the ‘genera’ allied to this are revised 
from the standpoint of comparative morphology. The features observed in detail are denticu- 
lation, hairs, internal celi walls, and sorus. Based on the observation of these features, the 
systems given by Copeland and Morton are critically discussed by the author who proposes 
several amendments for their systems, such as: Hymenophyllum s.str. is distinguished from 
Meringium only by the soral construction and Mecodium by the hairs and sorus; Hemicya- 


theon is identical with Meringium; and H. levingei is better segregated from Hymenophyllum 
s.str. 


The species belonging to Hymenophyllum s.1. were classified into 13 genera by 
Copeland (1938, 1947). Among them the ‘genera’ centering to Meringium are 
treated here to revise the system of the filmy ferns, According to the definition 
by Copeland, Meringium is the genus of some 60 species in the tropics and 
southern hemisphere, having a combination of such features as: 1) the segments 
and wings of axes denticulate at margin, 2) the involucre closed at the lower 
portion with bivalvate upper portion, 3) the receptacles growing indefinitely and 
extruded from the lips of involucre, 4) cell walls thicker and coarsely pitted. 
These characteristics are not completely represented by some species referred to 
Meringium. In this part of this series, several taxonomic characteristics of 
Meringium are revised, comparing with those of Hymenophyllum s.1., especially 
with the ‘genera’ allied to Meringium, e.g. Amphipterum, Buesia, Hemicyatheon, 
Leptocionium, Myriodon, and Hymenophyllum s.str. including H. levingei. 


Morton (1968) placed Meringium in Hymenophyllum subgen. Hymenophyllum 
and classified it as a distinct section named Ptychophyllum arranged next to sect. 
Hymenophyllum. In this system, he evaluated the denticulate margin of the 
ultimate segments, or of the wings as well, and separated Amphipterum and 
Leptocionium, placing them in subgen. Mecodium and Sphaerocionium, respec- 
tively. Copeland took much value, on the contrary, to the structure of sorus. 


In the following discussion all the species will be named under Hymenophyllum 
s.l., pending the nomenclatural discussion until the system of the filmy ferns will 
be revised as a whole. The names of subdivisions of the family will be adopted 
in accordance with Copeland’s system, or in some cases with Morton’s. 


Continued from the Fern Gazette 11 (2 & 3): 124. 


This study is partly supported by a grant in aid of scientific research of the Ministry 
of Education, Japan, no. 034047. 


63 


64 Gardens’ Bulletin, Singapore — XXX (1977) 


General features of Meringium sensu Copeland 


Before going further, the general features of Meringium will be summarized 
comparing with those found in the ‘genera’ allied to this, although some of them 
will be treated in detail in the next section. 


Rhizome. As usual in the case of Hymenophyllum s.1., the rhizome is long 
creeping, irregularly branched, wiry, nearly the same in thickness as or a little thicker 
than stipes. The apex is variously hairy with usually caducous hairs, the older 
portion being glabrous or very sparsely hairy. Roots are rather irregular in 
arrangement and bear hairs on all the surfaces. 


Fronds. The size of fronds is variable according to the species, or even within 
a species to some extent. The smallest is found in H. lobbii bearing fertile fronds 
less than 1 cm in length including stipes. On the contrary, the largest frond of 
H. penangianum at hand is nearly 35 cm in length. The fronds seem to elongate 
to some extent in humid mossy forest forming slender outlines. H. armstrongii 1s 
referred to Meringium and has only a few lobes, and H. lobbii, H. blandum and 
several others are pinnate in plan or at most with the pinnae a few times forked. 
The larger fronds are pinnate several times with simple or forked ultimate segments. 
The outline of the ultimate segments is long and slender, or comparatively short 
and broader downwards, 


Stipes are terete or winged nearly to the base. The hairs are present or absent 
according to the species; the presence on stipes correlates with that on other 
axes. The rachis is similar to the stipe, winged throughout in many cases. 
The wings on rachis and stipes are similar to the ultimate segments, denticulate 
or entire, plane or crisped, glabrous, and (2—) 4-12 (-25) cells broad. 


The ultimate segments which, as in the filmy ferns in general, consist of one 
cell layer except for the costae, are round to obtuse at apex, denticulate, or entire 
in some species, plane to distinctly crisped. 


Hairs. The hairs on the lower surface of axes, if any, are the same as those 
on rhizome. The coloration of the hairs is pale brown in the apical part of fronds 
and rhizome but dark brown in the older portion. The hairs are setose, pointed 
at apex, consisting of two to seven cells, and 0.5—2 mm in length. 


In H. johorense, H. reductum, and H. armstrongii, multicellular setae are 
found at margin of segments, and in Amphipterum the multicellular hairs are at 
margin of accessory wing. Except in these cases, no hairs are found on the laminar 
portion of the fronds in Meringium and its allies. 


Cell walls. The internal cell walls are thick and coarsely pitted in many species 
but not so thick and slightly waved in the others. This feature is evaluated by 
Copeland to distinguish Meringium from Hymenophyllum s.str. 


Sorus. The involucre is obconic at base with distinctly bivalvate lips. The 
size of involucre is rather variable and the lips are entire or denticulate. In most 
cases, it has been observed that the accessories on the involucre are like wings or 
bundles of hairs. The receptacles are clavate, grow indefinitely and extrude from 
the lips of involucre. 


Character phylogeny of four evaluated features 


Among the diagnostic features for defining Meringium and the ‘genera’ allied 
to this, four important characteristics will be treated here to revise the system of 
the species in question; They are denticulation, cell walls, hairs, and sorus. The 
evolutionary trends within each of these features will be discussed to elucidate 
the phylogenetic importance of them. | 


Meringium and allies | 65 


1. DENTICULATION 


The dentation of lobes is usually correlated with the veins for the megaphyllous 
leaves, though that of the filmy ferns in question is like that found in the lobes of 
the bryophytes without any actual relation to the veins. The marginal growth of 
the ultimate segments and wings as well is unequal, resulting in the irregular 
margin or the occurrence of denticulation, although the arrangement of it is regular 
in most cases (plate I). The denticulation here under consideration is, therefore, 
morphologically not homologous to that of vascular plants in general, though it 
will be described here under the term denticulation. 


The denticulation occurs in various forms in the species of Meringium. The 
wings of rachis and costae are entire or denticulate, and the denticulate segments 
and wings are either plane or crisped. The lips of involucre are toothed in many 
species and entire in the others. The occurrence of denticulation is also various 
according to the species. In H. edentulum the denticulation is rather rare, while 
H, meyenianum and allied species have regularly denticulate margin of segments, 
and the segments and wings of H. denticulatum are copiously denticulate and 
crisped to some extent. The last tendency proceeds comprehensively in H. 
acanthoides. 


The denticulate margin of ultimate segments is one of the key characters of 
Meringium, sharing it with Hymenophyllum s.str. Among the species of Meringium 
regarded by Copeland, however, there are several which have ultimate segments 
with entire margin: they are H. macroglossum, H. pachydermicum, H. penan- 
gianum, H. pollenianum, H. pulchrum, and H. ricciaefolium. Except for this 
characteristic feature of the entire margin of ultimate segments, they are close to 
the species of Meringium, or to H. meyenianum, in having the same morphology 
in hairs, cell walls, and sorus. 


Contrary to this case, a few species referred to Mecodium by Copeland are 
included by Morton in sect. Ptychophyllum, an equivalent of Meringium in the 
system of the latter: they are H. reinwardtii, H. samoense, H. taiwanense, and 
H. thuidium, In the diagnostic features other than the denticulate segments, these 
four species are different from H. meyenianum as noted by Copeland, and by 
Tagawa (1940) for the third species. The margin of ultimate segments is copiously 
crisped for these species, and the denticulation seems to be more irregular (Fig. 2). 
The arrangement of laminar cells at margin of segments is irregular in H. fimbria- 
tum, appearing somewhat denticulate, though the denticulation is different from 
that in Meringium species (Fig. 3). I prefer to exclude these species on the basis 
of their morphology in sorus, hairs, and cell walls in addition to the irregular 
denticulation at margin of segments, and would place them in Mecodium close to 
M. javanicum. 


Accessory wings and Amphipterum 


There are four species belonging to Amphipterum which is distinct in having 
accessory wings not in the plane of the lamina. Copeland (1938) distinguished 
Amphipterum generically from Meringium, although he (1937) had correctly 
suggested that ‘I would not consider it expedient to distinguish it generically or 
otherwise if the wings on the veins were the only distinction’. I would follow 
Copeland to consider that H. fuscum and three other species are close to each 
Other but are different from Meringium in combination of various features, 
admitting that the definition can be given solely upon the accessory wings on the 
veins. 


Accessory wings are prominent on veins, leaving no room to place any inter- 
mediate between presence and absence of this feature (Fig. 10). The cells compos- 
ing the accessory wings are quite the same as those of laminar surface in size, 
form, arrangement, and in structure of cell walls (Figs. 11 and 23). From this 


66 Gardens’ Bulletin, Singapore — XXX (1977) 


fact the accessory wings are considered to be the extra-outgrowths of the laminar 
surface in different levels from the usual ones. This kind of extra-expansion of 
laminar surface is unique to the filmy ferns but is not found in any other group 
of the megaphyllous plants. 


Except in the case of Amphipterum among the filmy ferns, the same kind of 
accessory wings is found in Dermatophlebium, or Hymenophyllum  subsect. 
Piumosa belonging to subgen. Sphaerocionium, although I know little of the 
eight species be!onging to this and am not able to conclude here whether they form 
a single taxon or not. In addition, the accessory wings are various in occurrence: 
three New Guinean species have the accessory wings on the upper side of veins as 
well, while the west Malesian H. fuscum has the wings only on the lower surface 
of veins; or among eight species of Sphaerocionium two species have the accessory 
wings only on the lower surface and six others have the wings on both the surfaces. 
In all the cases the cellular construction of the wings is identical with that of the 
ultimate segments of the species concerned. In the cases of both Dermatophlebium 
and Amphipterum, I have no sufficient materials at hand to conclude whether each 
of them forms a distinct taxon or not, though it will be safely said that the feature 
is derived parallel to each other in the above two distinct groups. 


‘Scales’ of H. levingei and Buesia 


H. levingei has hardly been recognized for a long time in spite of appropriate 
description and figures given by Clarke (1880). On the lower surface of midrib 
of ultimate segments are the ‘scales’ attached to the axes longitudinally in two 
rows and hardly continuous with the next ones on the same row but imbricating 
to the opposite ones on the neighbouring row. The base of ‘scale’ is 3-7 cells 
in breadth and upper half consists of several cells arranging in one row and 
appearing as an articulated hair with larger apical cell of clavate outline (Fig. 12). 
The ‘hairs’ on the upper surface are longer than the hair-like portion of the 
‘scales’ on lower surface, and they are articulated though not typically in construc- 
tion with thin side walls and thick septae (Fig. 13). The cellular construction of 
the ‘scales’ is similar to that of laminar portion of ultimate segments (Figs. 14 and 
15), so that the relationship between ‘scales’ and lamina is like that of accessory 
wing and Jamina in Amphipterum and others. Moreover, it will be noted here that 
the ‘scales’ and ‘hairs’ are in two rows on both the upper and lower surfaces of 
midrib of ultimate segments, and this arrangement is comparable with that of the 
accessory wings in Amphipterum. From these facts, it will be suggested that the 
‘scales’ and ‘hairs’ of H. levingei are the denticulation of the accessory wing, the 
laminar portion of such a wing being reduced in most cases. 


The morphology of the scales is common between 1H. levingei and the species 
belonging to Buesia. In the latter the ‘scales’ are often longer, with longer apical 
cells. The arrangement is typically in two rows, imbricate, as easily seen by the 
naked eye. 


Multi-directed projections and Myriodon 


I have once referred Bornean materials to Myriodon (Iwatsuki, 1968) based 
solely upon the literature for the characteristics of the latter, but am doubtful at 
present to regard them as Myriodon which has peculiar projections on various 
lines of the axes of various orders. In H. acanthoides the marginal teeth are in 
various directions owing to the comprehensive crispature of segments and the 
reduction of laminar portion, though the midribs, or axes, are still continuously 
winged in this species with the base of projections in one line. Contrary to this 
the projections on axes are recorded to be not in one line in the case of H. odonto- 
phyllum, differing in this respect from the Bornean materials. The reduction of 
laminar surface is not particular among the filmy ferns as known in Macroglena, 
Trichomanes setaceum and others. When I referred the Bornean materials to 


a 
4 


Meringium and allies 67 


Myriodon I considered that the materials are in the extreme form in the series of 
H. denticulatum — H. acanthoides — H. brassii, represented by the reduction of 
jaminar surface, though it is not probable if Copeland’s observation was correct 
concerning H. odontophyllum. 


The multi-directed projections of Myriodon may be speculated to have been 
resulted by the marginal denticulation and the splitting of accessory wing as 
schematically represented in Fig. 42. The teeth, or marginal projections, of segments 
and wings of H. acanthoides are similar to the projections of Myriodon and are 
also comparable with the ‘scales’ of H. levingei or of Buesia, which may be derived 
from the splitting of the accessory wing. In case the accessory wings are in double 
lines on both the surfaces, and the laminar parts as well as these accessory wings 
are split or reduced to remain as marginal teeth, the multi-directed projections will 
possibly be formed as in Myriodon, and the intermediate conditions are available 
as noted in the above description, although these ‘conditions’ do not necessarily 
represent the evolutionary intermediate but only the character phylogeny (Merkmals- 
phylogenie) is treated in this speculation. 


2. INTERNAL CELL WALLS (plate II) 


The internal walls of the laminar cells of Meringium are thick and 
coarsely pitted like those of Selenodesmium and some others belonging to Tricho- 
manes s.1, Contrary to these cases, the internal cell walls of Mecodium and 
Hymenophyllum s.str. are generally thin and straight or slightly waved at most, 
though there are species which do not accord well with the definition of this 
feature of the taxa concerned. Among the species which belong to Meringium, 
those with thinner walls are: H. armstrongii, H. blandum, H. bontocense, H. bivale, 
H., fejeense, H. macgillevirayi, H. multifidum, and H. viride. Even in these species, 
the cell walls are not straight but more or less waved. 


The thickness of the internal cell walls is also various among the species of 
Meringium, From the above observation, the difference in the structure of the cell 
walls is rather comparative, and not applicable to the difference in the taxa of 


higher rank, although a general tendency can be recognized as is treated such by 
various authors. 


The cell walls of Hymenophyllum s.str. are usually described as thin and 
straight, as illustrated by Copeland (1937) except for H. simonsianum. In my obser- 
vation, however, the cell walls are waved or pitted for all the species examined con- 
cerning Hymenophyllum s.str. The thickness is various according to the species, 
or even within the species for H. barbatum in which the southern form has thicker 
and more waved internal cell walls. 


Most of the species of Mecodium have thin and straight internal cell walls, 
although there are several species with thick and coarsely pitted cell walls, such 
as: H. crispato-alatum, H. exsertum, H. fimbriatum, H. flabellatum, H. javanicum, 
A. le ratii, H. montanum, H. oligosorum, H. opacum, H. riukiuense. The distribu- 
tion of the various forms of cell walls is similar to the case of Meringium and 
Hymenophyllum s.str., though the ratio of occurrence is different according to the 
‘genera’. From these facts, it is difficult to enumerate the structure of the cell walls 
as an important diagnostic feature to discriminate Meringium from the other groups 
belonging to Hymenophyllum s.1. 


The contents of cells are variously illustrated by van den Bosch (1861), 
though they have to be observed in the living condition to evaluate the taxonomic 
significance. In this study the living materials were available only for a few species. 


68 Gardens’ Bulletin, Singapore — XXX (1977) 


3. HAIRS ON RHIZOME AND FROND 


The so-called articulated hairs are found on the upper surface of the axes in 
H. levingei and in Buesia, though this kind of ‘hairs’ is observed as to be identical 
with the ‘scales’ on the lower surface. The discussion on such ‘hairs’ has been given 
in the section on denticulation and is not appropriate to note in the section on 
hairs, | 


At the margin of the accessory wing on the veins in Amphipterum, there are 
the hairs identical with those found on the axes and rhizome of most species of 
Meringium and Amphipterum, The general morphology of this type of hairs is 
described in the foregoing pages in the section of general morphology. The basal 
cell of the hairs is attached to the axes or marginal cells of the accessory wing at 
the base or at the middle portion (Fig. 35) and the hairs are adpressed. Hairs 
of this type are also found in the species of Hemicyatheon, Hymenophyllum s.str., 
Leptocionium, and several species of Mecodium. In these species as well as in 
Meringium, the hairs are restricted to the axis and never observed at the margin 
of lobes. In this respect it is rather peculiar to observe the hairs at the margin 
of the accessory wing in Amphipterum. In some species of Meringium, the fronds 
are nearly glabrous, or the hairs are restricted to the very young portion of fronds 
and rhizome, though the structure of the hairs if any is the same in all the species 
belonging to the ‘genera’ cited above. 


On the veins or especially at their junction, most species of Mecodium 
bear rather sparsely, smal! multicellular hairs consisting of several sub-transparent 
cells with thin walls (Fig. 39). This kind of hairs is quite different from the above 
described hairs found in Meringium and ‘genera’ allied to this. The stellate hairs 
of Sphaerocionium (Fig. 36) were observed in detail by Morton (1947) who 
applied the distribution of hairs to the subdivision of that ‘section’. The stellate 
hairs are only known in Sphaerocionium, though the hairs of some species of 
Microtrichomanes were referred to the former hairs (Iwatsuki, 1975). 


Marginal setae of H. johorense, H. reductum, and H. armstrongii 


The morphology of H. johorense was well described by Holttum (1929, 1955) 
and Copeland (1937), although the marginal setae were illustrated only by the 
line drawing. The marginal setae are in two to six cells, with oblique septa giving 
hooked appearance of setae, dark brown, polished, with thick walls and pointed 
apex (Figs. 37-38). The setae are similar to those of Trichomanes digitatum group 
in appearance except for the multicellular construction in contrast with the 


unicellular setae of the latter. Similar setae are also found in Didymoglossum which 
belongs to Trichomanes s.1, 


The fronds of H. johorense are small, simple to 5-6 lobed, branched nearly 
dichotomously, comparable to Trichomanes digitatum in this frond form, though 
this is also known in various dwarfed species as noted in the first part of this 
series. The margin of lobes is entire in H. johorense, not specialized except for 
bearing setae, the cell walls are thick and coarsely pitted, and the receptacles are 
extruded. We have no species of Meringium comparable to this species except for 
H., reductum and the systematic position of them is unknown at present. Copeland 
considered H. johorense as a member of Microtrichomanes but transferred H. 
reductum to Meringium without any reasonable interpretation. 


H. armstrongii is a small fern in New Zealand, having the fronds simple to 
four-lobed, arranged subdichotomously or flabellately. Morton treated this species 
as a second member of subgen. Craspedophyllum, and his treatise seems to be 
probable considering only thin cell walls and deeply cleft bivalvate involucre, 


Meringium and allies 69 


though the presence of marginal setae makes it doubtful to this conclusion. At 
margin of the ultimate segments of H. armstrongii are the setae of the same type 
as those of H. johorense. The marginal cells of the lobes are polished dark-brown, 
similar to the cell walls of the setae. In Craspedophyllum, however, the cells of a 
marginal row of the lobes are specialized without any marginal setae. The marginal 
cells of H. armstrongii are not specialized in structure, differing only in the dark- 
brown coloration. Anyway, it will be advisable at present to exclude H. johorense, 
H. reductum, and H. armstrongii from Meringium on the basis not only of the 
marginal setae but also of various other features. 


4. Sorus 


The position of sorus is paratactic in all the species in question. The involucre 
is obconic in the lower half with bivalvate upper portion. In this diagnosis the 
involucre is similar to that of Crepidomanes especially when pl. 16 of Copeland 
(1937) is compared with pl. 27 of Copeland (1933) as an example. The involucre 
of Crepidomanes is usually longer as a whole with longer tube and entire lips 
usually having the pseudoveins. In many cases the lips of involucre are denticulate 
at margin in Meringium, but the denticulation at lips of involucre is not in 
accordance with the denticulation at margin of segments and wings. On the surface 
of involucre, usually on the lower obconic part, there are the accessory projections 
in some species, especially in H. denticulatum, H. acanthoides and others. The 
depth of cleft is variable to some extent, and the obconic portion is either narrowly 
winged or not. 


I have made a preliminary observation on the development of sorus in H. 
polyanthos and found that at first a small cup-shaped involucre is developed 
which becomes deeply cleft at maturity. Further observation is necessary to 
elucidate the character phylogeny of sorus. 


The receptacles are clavate to cylindrical, growing indefinitely, and extruded 
from the lips of involucre. The form of receptacles is variable among the species 
of Mecodium, including capitate and clavate receptacles and longer involucre, but 
never extruded. The extrusion of the receptacles is various according to the species, 
although it will be notable that the exserted receptacles are found only for the 
sorus with obconic base. In Mecodium and Hymenophyllum s.str., the involucre is 
cleft nearly to the base forming no obconic basal portion, and the receptacles are 
never extruded even in the case when they are clavate in structure. 


Hymenophyllum s.str. is discriminated from Meringium by the structure of 
sori, though the distinction is in some cases obscure. I have examined only a small 
number of specimens of American species, although I can point out that several 
species which are referred to Meringium have deeply cleft bivalvate involucre 
which forms a tube only at the basal portion. New Caledonian H. dimidiatum was 
included in Meringium by Copeland, and this is followed by Morton, as an isolated 
species, even with the description ‘receptacle, so far as seen, included’. Copeland 
noted that ‘sori .... cleft about half-way down’, though in fact the sori are deeply 
cleft nearly to the base and form obconic base when the sori are rather deeply 
placed at apex of the ultimate segments. From these facts, H. dimidiatum seems 
to be better placed in Hymenophyllum s.str. Contrary to that case, Australian H. 
cupressiforme is said to belong to Hymenophyllum s.str., by both of the above 
authors, though the receptacles are clavate and are larger, extruding in some cases 
from the lips of involucre. The small extrusion seems to depend upon the larger 
size of clavate receptacles, different from the long extruded receptacles typical 
for Meringium. 


70 Gardens’ Bulletin, Singapore — XXX (1977) 


Classification of Meringium and ‘genera’ allied to this 


It is recommended that all the species ever described be examined before 
proposing a system in the categories lower than family. In this study, however, the 
observation was made only for some representative species, pending the detailed 
comparison for every species concerned. The discussion on the system is, therefore, 
possible only in a limited sense, and a few comments will be made on the ever 
proposed systems. 


Before the discussion on the relationship among the species of Meringium 
group, the systems proposed by Copeland and Morton will be summarized in a 
table: 


Copeland (1947) 
I. Gen. 1 Mecodium, 2 Craspedophyllum, 3 Hemicyatheon 
II. Gen. 4 Sphaerocionium, 5 Apteropteris, 6 Microtrichomanes 
UI. Gen. 7 Hymenophyllum 


IV. Gen. 8 Meringium, 9 Amphipterum, 10 Myriodon, 11 Buesia, 12 Leptocionium, 
13 Rosenstockia. 


Morton (1968) 
Gen. HI. Rosenstockia 
Gen. IV. Hymenophyllum 
Subgen. | Hymenophyllum: Sect. 1 Hymenophyllum, 2 Buesia, 3 Ptychophyllum, 
4 Eupectinum, 5 Myriodon 
Subgen. 2 Sphaerocionium: Sect. 6 Sphaerocionium (Subsects. Ciliata, Plumosa, 
Hirsuta, and Leptocionium), 7 A pteropteris 
Subgen. 3 Craspedophyllum: Sect. 8 Craspedophyllum 
Subgen. 4 Hemicyatheon: Sect. 9 Hemicyatheon 


Subgen. 5 Mecodium: Sect. 10 Mecodium (Subsects. Mecodium, Amphipterum, and 
Diplophyllum). 


1. MERINGIUM AND HYMENOPHYLLUM S.STR. 


In definition Hymenophyllum s.str. is distinguished from Meringium ‘by the 
more deeply cleft involucre, the shorter receptacle, the absence of peculiarly 
(pitted) thickened cell wall, and usually smaller size’ (Copeland, 1838: p. 38). 
There are definitions by various authors. but the above seems to be a representative 
figuration. Morton cites the direction of sori for Hymenophyllum s.str., but this is 
true for only a few number of species. 


Copeland (1938) enumerated 14 species for Hymenophyllum s.str. by his 
definition. One of them was transferred to sect. Ptychophyllum by Morton (1968) 
who added 11 species to sect. Hymenophyllum mostly following the suggestion 
given in the Index of Copeland (1938). I can not say exactly at present how many 
species belong to Hymenophyllum s.str., which may be classified into two: the 
group of H. peltatum represented by entire involucral lips include H. antarcticum, 
H. perfissum, H. subdimidiatum, and H. wilsonii; the other is the group of H. 
thunbridgense with denticulate involucral lips, including H. barbatum, H. dimi- 
diatum, H. revolutum, and H. simonsianum. H. cupressiforme seems to belong to - 
the former, though the lips are slightly dentate with shortly extruded receptacles, 
deeply cleft involucre, and thin but pitted internal cell walls. I have no sufficient 
knowledge on the southern species, although Copeland pointed out that the 
dwarfed species in the Far South, e.g. H. minimum, H. pumilo, and H. pumilum, 
were the representatives which were difficult to be distinguished between Meringium 
and Hymenophyllum s.str., the former two being actually included in sect. 
Ptychophyllum by Morton. 


Meringium and allies 71 


H. levingei has by far no direct relationship to H. thunbridgense. In having 
the ‘trichomanes’ on the axes of fronds, this resembles Buesia, though this particular 
feature may have been evolved independently in H. levingei and in Buesia. H. 
levingei is similar to Buesia in this feature but has entire margin of ultimate 
segments and deeply cleft involucre with clavate but not extruded receptacles. This 
species should be segregated from Hymenophyllum s.str., probably at the same level 
as Buesia from Meringium. 


The cell walls are not distinctly different between Meringium and Hymeno- 
phyllum s.str., and the pale brown multicellular hairs are found in both of them. 
The sole feature to discriminate these two is the morphology of sorus, especially 
in the obconic base of involucre and extruded receptacles in the former, From 
these facts, Hymenophyllum s.str. and Meringium are considered to be close to each 
other accepting the system of Morton (1968) at least concerning this part. 


2. MERINGIUM AND MECODIUM 


Morton included in sect. Ptychophyllum several species which were included 
in Mecodium by Copeland, e.g. H. reinwardtii, H. samoense, H. taiwanense, and 
H. thuidium. The interpretation in the structure of leaf margin was different between 
the above two authors. As noted in the previous pages in the section of denticula- 
tion, I prefer to follow Copeland keeping the above four species in Mecodium. 


Among the Asiatic species of Mecodium, those of the group of H. javanicum 
is represented by waved or crisped margin of the ultimate segments and wings, 
thus representing the appearance similar to Meringium. Some of the species 
belonging to this group, e.g. H. fimbriatum, H. javanicum, and H, riukiuense, have 
denticulate lips of involucre and pitted internal cell walls. In these features they 
share the same characteristics with Meringium, though they appear to be identical 
as the result of the evolution along different courses. 


H. macroglossum and H. pachydermicum were placed by Morton in sect. 
Sphaerocionium subsect. Ciliata ‘without undue strain’, contrary to the treatise of 
Copeland who included them in Meringium. As the hairs are quite different between 
Sphaerocionium and the other ‘genera’ belonging to Hymenophyllum s.1., we 
cannot accept the transferance by Morton as appropriate. We would refer to the 
fact that the above two species are close to some species with dense hairs belonging 
to Mecodium, e.g. H. exsertum, H. gardneri, and H. oligosorum, in hairiness, 
structure of internal cell walls, entire margin of segments, texture, coloration, and 
pinnation with broader wings giving the general appearance of shallow incision, 
but different only in the structure of sorus. The difference in soral construction is 
considered in this paper to record exactly the phylogeny among the species of the 
filmy ferns, though the pale brown multicellular hairs are common in Meringium 
and Hymenophyllum s.str. but not found in Mecodium except for the above species. 
H, barbatum, a representative of Hymenophyllum s.str.. is also similar in appea- 
rance to the above mentioned species except for the denticulation at margin of 
segments and not so thick cell walls. 


Morton noted that only four species of Meringium in the sense of Copeland 
had segments with entire margin but in fact the latter author included four more 
species without denticulate segment in the latter, and Morton enumerated these 
species in sect. Ptychophyllum. As noted in the section of denticulation, the entire 
margin is rather peculiar among the species of Meringium, but H. penangianum 
seems to be close to H. holochilum except for the entire margin of segments. H. 
pachydermicum and H. macroglossum are different in various features from H. 
meyenianum, but still belong to Meringium in the broader sense. 


2 Gardens’ Bulletin, Singapore — XXX (1977) 


Morton distinguished Ptychophyllum and Mecodium in the level of subgenus, 
based chiefly on the structure of the margin of segment. In addition to the 
difference in denticulation, we can list up the difference in the structure of sorus 
and hairs between Meringium and Mecodium, admitting that the difference between 
them is comparatively smaller than the one between these two and Sphaerocionium. 


3. THE ‘GENERA’ ALLIED TO MERINGIUM 


Copeland gave a diagram of affinity of genera and showed that the close 
allies to Meringium were Amphipterum, Buesia, Leptocionium, and Myriodon, 
adding to them Hemicyatheon and Rosenstockia of doubtful relationship. In the 
opinion of Morton, on the contrary, the closest allies to Meringium are Buesia, 
Hymenophyllum s.str., Myriodon and a small group in South America named 
Eupectinum. He lowered the status of Leptocionium and Amphipterum to sub- 
sections under the subgenera Sphaerocionium and Mecodium respectively, that of 
Hemicyatheon to a distinct subgenus but retained Rosenstockia as a separate 
genus. Thus in his classification of the Hymenophyllaceae he admitted six genera. 


Amphipterum. Morton considered that, only two species among four 
enumerated by Copeland, belonged to this taxon separating the other two on the 
basis of having the segmental margin denticulate. The systematic evaluation on 
denticulation has repeatedly been discussed in this paper which concludes that we 
can not distinguish any taxa of the filmy ferns in question based solely on this 
feature. I found no sound reason in the treatise of Morton and consider four, or 
three, species treated by Copeland are close to each other. 


Thus confined, Amphipterum includes the species having the leaf margin 
denticulate or entire, although the accessory wing on veins not in the plane of the 
lamina is common to the species in question. The structure of sorus, hairs, internal 
cell walls, and the general appearance are not different between Meringium and 
Amphipterum, and it will be accepted here to keep Amphipterum as a distinct 
taxon very close to Meringium. 


Buesia, Five species were referred to this by Morton (1968) who noted that 
he was uncertain that it even needed to be distinguished as a section. As was 
suggested by him, Buesia should better be compared with the species having the 
accessory wing, according to the speculation that the ‘scales’ may be the broken 
accessory wing. 

Myriodon. The multi-directed projections of Myriodon were inferred as having 
been resulted from the occurrence of laminae in more than two planes and the 
splitting of laminar surface or reduction of lamina, remaining only the teeth. 
According to the above interpretation, that feature is particular, but Myriodon is 
quite similar to Meringium except for the above mentioned feature. Myriodon is 
accepted here as a monotypic taxon very close to Meringium, the conclusion 
being identical with that given by Copeland and by Morton. 


Leptocionium. Indicated by the morphology of setae, this is considered as a 
relative of Sphaerocionium but not so close to Meringium. In this I prefer to follow 
not Copeland but Morton. 


Hemicyatheon. This is recognized by Copeland on the basis of two species 
which were considered by him closely related to each other but had fallen into 
Meringium and Mecodium by definition, respectively. Morton maintained Hemi- 
cyatheon as a subgenus based on the type species, referring another species to 
subgen. Hymenophyllum. 


H. deplanchei is not very close to H. meyenianum, though there is no problem 
to include the former in Meringium, especially by such features as denticulate 
margin of ultimate segments, thick and pitted internal cell walls, obconic involucre 


Meringium and allies 73 


with denticulate lips cleft to the half-way, and exserted receptacles. Copeland’s 
negotiation to place H. deplanchei in Meringium was based on his speculation that 
this species was close to H. bailayanum which was different from Meringium in 
entire margin of ultimate segments, although we refuse to separate any species from 
Meringium based solely on this feature. Judging from the soral structure and 
morphology of cell walls, H. bailayanum seems to belong to Meringium in broader 
sense, probably next to H. deplanchei, with the result again that the entire segment 
is not so particular to discriminate the taxa in a rank higher than species. Both 
Copeland and Morton consider that H. bailayanum is close to Mecodium and 
distinct from Meringium based on the entire margin of segments, although it is 
similar to Meringium in soral structure and cell walls which actually indicate 
the relationship among the species of the filmy ferns in question. Thus the two 
species included in Hemicyatheon by Copeland may better be transferred to 
Meringium. 

Rosenstockia. I have little information to add to this monotypic ‘genus’, 
pending the further study when fresh materials are available. 


4. SUBDIVISION OF MERINGIUM 


There are several species groups separable from H. meyenianum and its close 
allies, although it is rather difficult to diagnose the groups within a few words. 
Moreover, there are several species which are included in Meringium by Copeland 
but are to be separated from this. 


H. johorense, H. reductum, and H. armstrongii are distinct from Meringium 
in having dark brown marginal setae. As noted in the section of hairs, these three 
are distinct from the other members of Meringium. H. johorense and H. reductum 
form a small taxon not so far from Microtrichomanes. I have little information 
to add at present to H. armstrongii pending further investigation. It is doubtful 
that the latter is referable to Craspedophyllum as done by Morton. 


Both H. lobbii and H. blandum are dwarfed species and apparently similar to 
each other, although it is difficult to know whether their resemblance is the result 
of parallel evolution or of the phylogenetic similarity. It is possible to recognize 
the similarity between these dwarf forms and the larger species without any actual 
evidence. [ am sure that these species are not close to H. johorense and H. reductum 
irrespective of their apparent similarity. 


H. pachydermicum and H. macroglossum form a group distinct from the 
others in having dense hairs on axes beneath, broader wings of axes and short 
entire segments which are broader towards base, giving an appearance of fronds 
less dissected, and brownish in dried condition. I have not seen actually the 
materials of H. pulchrum which seems to belong here by description and figure 
given by Copeland. 

Many of the Southeast Asian species referred to Meringium are close to H. 
meyenianum having the characteristics described for typical Meringium. I am not 
sure at present whether the group of H. holochilum is separable from H. 
meyenianum group or not. H. penangianum and H. edentulum with entire or 
subentire margin of ultimate segments belong to the group of H. meyenianum in 
very strict sense. 


I know little at present about the American species, and the southern species 
are still not observed in the living condition. H. deplanchei and H. bailayanum 
form a separate group even when they are included in Meringium, and no close 
allies can be represented here. 


74 Gardens’ Bulletin, Singapore — XXX (1977) 


5. CONCLUSIVE REMARK 


It is necessary to make a detailed revision of the species concerned before 
having a systematic conclusion of Hymenophyllum s.1., although it will be still useful 
to give here a system rather diagrammatically summarizing the above discussion for 
the basis of further investigation. This is a tentative scheme and is a modification 
of the systems given by Copeland and by Morton. 


Hymenophyllum s.1. 
Hymenophyllum 


Hymenophyllum s.str. (Hymenophyllum s.str. with two subgroups, H. levingei, 
and Eupectinum) 


Meringium (Meringium s.str. with several subgroups, Amphipterum, Buesia, and 
Myriodon) 


Mecodium 
Sphaerocionium 


References 


van den Bosch, R. 1861. Hymenophyllaceae javanicae. Verh. kon Acad. Wetensch. 
Amsterdam [X-—6: 1-67. 


Clarke, C. B. 1880. A review of the ferns of Northern India. Tr. Linn, Soc. II. 
Bot, b. 425-61. 


Copeland, E. B. 1933. Trichomanes. Phil. J. Sci. 51: 119-280. 
1937. Hymenophyllum, Phil. J. Sci. 64: 1-188. 

1938. Genera Hymenophyllacearum. Phil. J. Sci. 1-110. 
———— 1947. Genera Filicum. Waltham, Mass. 247 pp. 


Holttum, R. E. 1929, New species of ferns from the Malay Peninsula. Gard. Bull. 
S.S. 4: 408-410. 


1955. A revised flora of Malaya II. Ferns of Malaya. Singapore. 643 pp. 


[watsuki, K. 1968. Contributions to the classification of the filmy ferns (2). Acta 
Phytotax. Geobot, 23: 117-125, in Japanese. 


——-— 1975. Studies in the systematics of filmy ferns I. A note on the identity 
of Microtrichomanes. Fern Gaz. 11 (2 & 3): 115-124. 


Morton, C. V. 1947. The American species of Hymenophyllum section Sphaero- 
cionium, Contr. U.S. Nat. Herb. 29: 139-201. 


1968. The genera, subgenera, and sections of the Hymenophyllaceae. 
Contr. U.S. Nat. Herb. 38: 153-214. 


Tagawa, M. 1940. Studies on Formosan ferns 2, Acta Phytotax. Geobot. 9: 139-148. 


Plate I. Denticulation. Figs. 19. Margin of ultimate segment, x 100: Fig. 1, H. poly- 
anthos (Iwatsuki et al. M 13034); Fig. 2, H. taiwanense (type); Fig. 3, H. fimbriatum (Tagawa 
3347); Fig. 4, H. barbatum (Tagawa et al. T 1502); Fig. 5, H. cupressiforme (Muir 765): 
Fig. 6, H. penangianum (Mizutani 252245); Fig. 7, H. meyenianum (Iwatsuki et al. M 13549); 
Fig. 8, H. denticulatum (Iwatsuki et al. T 14579 bis); Fig. 9, H. acanthoides (Iwatsuki et al. 
M 13235). Figs. 10-11. Accessory wing of H. fuscum (Iwatsuki et al. S 1063): Fig. 10, 
ultimate segment with accessory wing, x 100; Fig. 11, Accessory wing, x 400, compare 
with II-23. Figs. 12-15. ‘Scales’ and ‘hairs’ of H. levingei (Kanai et al. 8249), x 100: Fig. 12, 
‘scales’ on lower surface; Fig. 13, ‘hairs’ on upper surface; Figs. 14 & 15, comparison of 
‘scales’ and segment, with focus on ‘scales’ in 14 and on segment in 15. 


Plate II. Internal cell walls, all x 400: Fig. 16, H. meyenianum (Iwatsuki et al. 
M 13549); Fig. 17, H. denticulatum (Iwatsuki et al. T 14579 bis); Fig. 18, H. blandum 
(Iwatsuki ei al. M 13515); Fig. 19, H. multifidum (Pichi Sermolli 6257); 20, A. bontocense 
(Tagawa et al. T 4815); Fig. 21, H. johorense (Mizutani 2724 a); Fig. 22, H. deplanchei 
(Balansa 1641); Fig. 23, H. fuscum (Iwatsuki et al. S 1063); Fig. 24, H. barbatum (Tagawa 
et al. T 1502); Fig. 25, H. simonsianum (Kanai et al. 725175); Fig. 26, H. wilsonii (Jermy 
et al. 8473); Fig. 27, H. cupressiforme (Muir 765); Fig. 28, H. levingei (Kanai et al. 8249); 
Fig. 29, H. polyanthos (Iwatsuki et al. M 13034); Fig. 30, H. fimbriatum (Tagawa 3347); 
Fig. 31, H. oligosorum (Iwatsuki 6657); Fig. 32, H. exsertum (Kanai et al. 725170); Fig. 33, 
H. hirsutum (Standley 21479). 


Plate III. Hairs and denticulation. Figs. 34-39. Hairs: Fig. 34, multicellular hair of 
H. exsertum (Kanai et al. 725170, x 100; Fig. 35, multicellular hair at margin of accessory 
wing of H. fuscum (Iwatsuki et al. S 1063), x 100; Fig. 36, stellate hairs of H. hirsutum 
Standley 21479), x 100; Fig. 37, marginal setae of H. johorense (Mizutani 2724 a), x 100; 
Fig. 38, oblique septa of Fig. 37, x 400; Fig. 39, multicellular hair of H. polyanthos 
(Iwatsuki et al. M 13034), x 400. Figs. 40-41. H. acanthoides (Iwatsuki et al. M 13235): 
Fig. 40, a plant, x 5; Fig. 41, segment with crisped denticulation, x 100. 


‘gq {quowses eunjjn jo uonenonusp ‘D ‘suIM 
AuasojAyd IajOvIVYS & JOJ SWTDYSs [eONOYIOdAH ‘Sp “314 ‘uinuissisojid *"H 
-OYIT “pp “BI ‘unjnqging *H “tv ‘BIg ‘soyjundjod "HY ‘Zp “Bt :snios J 


8¥ 


a 
i 


Cece eet FP eeeteaeeces 
te ee ee 


eeeccsse gece Se Perens 


eeceeer eessteeer eames 


NS 
oO 


PPO EP SEPOCEO EPO EHOE 
SOC Cee eeraeseee eraser eee 
eQbeececess coeeesere® 


Cr tec ece-- — er 


| ‘U0polApPp ‘g 
‘pisang ‘9 ‘wumi8uuapw ‘L 9 p “1asuiaay “HH ‘¢ “uinsaidiydmp ‘¢ 2% 7 “wnipooap “| -4}99} 9} AjUO 3uluIeWol UONIod IeuIWe, Jo UoTONpel 
eur Aiossoo0" jo sunyyids ‘g ‘8uim Arossos9v Jo JUoudo[sAap ‘Pp :UOTe[NOWUSp oY} SuTUIEDUOS 
‘Lp “B1q ‘lasoy “H ‘9p “S14 “uouvjnoyuap “H 
O SSUIMPIP SUIT ‘Lp-Zp ‘S3IJ ‘UONe[NOWUep pue snIog A] ed 


-- 
- 
aaannse° 

- 
Swern ce o°"” 


e 


Sh ‘SI ‘wniwsip saupwu 


44 


Calamus caesius and Calamus trachycoleus Compared 
by 
J. DRANSFIELD 


Royal Botanic Gardens, Kew 


Summary 


Two closely related rattan palms, Calamus caesius and C. trachycoleus are shown to have 
remarkably different habits; the silvicultural significance of the difference in habit is discussed. 


Throughout the Lower Barito area of South Kalimantan, Indonesian Borneo, 
a rattan species receives intensive cultivation. This species, Calamus trachycoleus, 
is the only species cultivated on a large scale (i.e. thousands of hectares). It is 
closely related to Calamus caesius which also receives some degree of cultivation 
in the area and elsewhere. Though so similar to Calamus caesius, C. trachycoleus 
differs in one feature of its growth form; this difference has a major effect on the 
habit of the rattan and is one of the most important features of the species which 
has led to its ultimate supremacy as a plantation rattan. Holttum (1955) in his 
classic paper on growth forms of monocotyledons has indicated the importance 
of an understanding of the basic growth forms of monocotyledons; here is an 
example from the rattans where a difference in the degree of growth of one feature 
of a basically similar growth pattern differentiates between two species and 
accounts for the economic success of one over the other in cultivation. 


DESCRIPTION OF THE GROWTH FORMS OF CALAMUS CAESIUS AND 
C. TRACHYCOLEUS 


CALAMUS CAESIUS BLUME 


This rattan is very widespread being found in Sumatra, the Malay Peninsula 
including Southern Thailand, and Borneo, at altitudes ranging from near sea-level 
in coastal peat swamp forest to 1000 m or more in the hills; the wide range may 
in part be due to the fact that it receives a rudimentary cultivation and is some- 
times planted in primary forest near villages (Dransfield pers. obs. and hearsay 
from villagers in Borneo). It also seems to tolerate a great range of soil conditions 
from seasonally flooded alluvial clay soils, peat-swamp soils to well-drained steep 
soils on varying substrata in Hill Dipterocarp Forest. It is however commonest 
as a lowland plant on alluvial flats beside rivers. Throughout Borneo, the indigenous 
people often plant a few clumps of this and occasionally other species at the edge 
of their villages or near their longhouses. Calamus caesius without doubt produces 
the best quality small diameter class (7-15 mm) cane of all rattan species entering 
the rattan trade. 


Calamus caesius produces rather dense clumps of many aerial stems, often 
more than 15 in number radiating from a condensed system of short underground 
rhizomes. Initial growth of the seedling produces an orthotropic stem about 1 cm 
in diameter including the leafsheaths. This seedling stem branches from the basal 
nodes to produce one to three sucker shoots which are also orthotropic; subsequent 
suckering from these sucker shoots produces short subterranean rhizomes c 3 cm 
in diameter and up to 10 cm long which bear pale brown scale leaves and short 
condensed internodes. Such rhizomes eventually (growth rates of the rhizomes 
are not known) metamorphose into rapidly growing orthotropic shoots of diameter 
1.5 — 2.5 cm including the leaf-sheaths with lower internodes 50-100 cm or more 


75 


76 Gardens’ Bulletin, Singapore — X XX (1977) 


in length bearing at the base leaves with highly reduced laminae often consisting 
of rachis alone and long-sheathing bases. Subsequent leaves develop more and 
more leaflets until the adult leaf shape is attained with the rachis bearing 10 or 
more grouped leaflets on each side, the rachis tip terminating in a cirrus. As 
the rhizome metamorphoses into an orthotropic stem, the two nodes at the area 
of metamorphosis each develop a branch with the potential of continued rhizoma- 
tous growth though the potential may not always be realised. Some of the 
branches remain dormant as bulb-like shoots. As each rhizome grows into an 
aerial stem, there is the potential for replacement by two new rhizomes and hence 
exponential increase in the number of aerial stems in the clump. However, because 
of the shortness of the rhizomes, many of the potential new rhizomes become 
juxtaposed and squashed and in this condition remain as bulb-like dormant shoots 
as described above. Further development of these dormant shoots depends on the 
opening up of the clump either effected by death of orthotropic shoots or by 
clearance by man of debris during cultivation of this species (pers. comm. from 
villagers in Borneo). 


CALAMUS TRACHYCOLEUS BECC. 


This rattan is only known from South Borneo within the watersheds of the 
rivers flowing into the Java Sea — i.e. the Barito, Kapuas (Kalimantan Tengah), 
Kahayan, Mendawai, Sampit and Seruyan Rivers. In this area it is found growing 
on seasonally-flooded riverbanks on alluvial clays and the margins of swamp forest. 
It receives intensive cultivation in the Barito Selatan area upriver from Kuala 


— 


fais Wy 
a 


nN 
SP, o? ya 
J 


4 me My if 
fha> WZ 
Ay itr fos 
Y, EH No aa 
lt Ail, 


y 


1. The base of a clump of Calamus caesius with most roots removed; note the short 
rhizomes, bulb-like dormant shoots, and the smaller diameter of the aerial shoots com- 
pared with that of the rhizomes, (Dransfield 3933, Sungei Jaya, Kalimantan Tengah, 
4.2.1974) 


Calamus 


2. A piece of a stolon (“selantar’) of 


Calamus trachycoleus at the point of 
metamorphosis into an orthotropic; stem 
sheaths have been removed to show the 
production of two branches at the point 
of metamorphosis. Dransfield 3929, 
Sungei Jaya, Kalimantan Tengah 
4.2.1974). 


77 


Kapuas on the Barito River, being grown 
in plantations on riverside seasonally- 
flooded alluvia] soils. Though eminently 
successful as a plantation crop and in the 
wild, apparently, as a riverbank colonizer, 
it is unknown elsewhere in Borneo. 
Calamus trachycoleus produces a small 
diameter class cane (7-15 mm) with 
internodes generally shorter than those of 
Calamus caesius and of not quite such 
good quality; yet it accounts for about 80 
per cent of the rattan trade on the Barito 
River. 


Calamus trachycoleus is immediately 
distinguishable from C, caesius in produc- 
ing diffuse open colonies rather than dense 
clumps; rather than having a condensed 
system of short underground rhizomes, it 
spreads by means of lax above-ground 
stolons. Initial growth of the seedling is 
similar to that of C. caesius, the first stem 
and one or two suckers being orthotropic. 
Subsequent branching results in robust 
stolons up to 4 cm in diameter which grow 
along the soil surface or slightly raised 
above ground level on short adventitious 
roots. The stolons (known locally as 
“selantar’) bear sheathing pale brown 
scale leaves and short internodes 4-7 cm 
long and like the rhizome of C. caesius 
metamorphose into orthotropic stems. The 
length of the stolons from point of origin 
to point of metamorphosis may be 3 m 
or more. As in Calamus caesius, two 
branches are produced at the area of 
metamorphosis, one each from adjacent 
nodes, but unlike C. caesius, both branches 
grow out to produce new stolons unless 
damaged; no dormant branches have been 
observed and this is regarded as being 
correlated with the open growth of the 
clump and apparent lack of competition 
between the branches. 


Each branch in Calamus trachycoleus 
is apparently adnate to the internodes 
of the proceeding leaf and is hence carri- 
ed out of the scaleleaf axil; however their 
position suggests an axillary origin rather 
than an internodal or other anomalous 
origin. This feature of Calamus trachyco- 
leus deserves anatomical investigation. Of 
the two branches produced at each meta- 
morphosis, the proximal tends to develop 
before the distal and may be already 30 cm 
in length before the distal emerges 


78 Gardens’ Bulletin, Singapore — X XX (1977) 


from the scale leaves, Because of distortion the proximal branch of the stolon 
appears to continue the growth of the subtending stolon and at first sight the 
orthotropic stem appears as the branch rather than the main axis. If young material 
is examined at a stage when a stolon tip begins to grow upwards, the two new 
stolons are seen to be branches rather than one of them being a continuation of the 
stolon. 


Unlike Calamus caesius, C. trachycoleus is a rapidly invasive species and the 
potential of exponential increase in number of aerial stems is usually realised, Once 
a plantation of C. trachycoleus has been established very little cultivation is neces- 
sary and harvests can be made after an initial 7-10 year period at 2 yearly intervals. 
In C. caesius on the other hand, clumps require clearance of debris to encourage 
development of new shoots and in the Barito Selatan area of South Borneo at any 
rate, only two main harvests are obtained, an initial one after 7-10 years followed 
by a second after a further four years; after the second harvest the clumps are 
supposedly exhausted. 


DISCUSSION 


In his original description of Calamus trachycoleus Beccari (1913) comments 
that the rattan is allied to C. caesius but most closely related to C. pogonacanthus. 
Recent fieldwork in Borneo has allowed the present author to make several 
collections of C. pogonacanthus which is now seen to belong to a distinct group of 
species in Calamus all with the peculiar feature of bearing both a cirrus and a 
flagellum, Cirrus (barbed whip at the end of the leaf representing an extension 
of the leaf rachis) and flagellum (barbed whip borne on the leaf sheath and 
representing a sterile inflorescence) are the two major climbing organs found in 
Malesian rattans and are usually mutually exclusive. The presence of both in 
C. pogonacanthus and a few other species is hence noteworthy. C. trachycoleus 
bears a cirrus only and hence is here regarded as being more closely related to 
C. caesius than to C. pogonacanthus. Furthermore, in the field, without reference 
to the remarkable difference in habit, C. caesius and C. trachycoleus are distinguished 
with some difficulty — both species are about the same size, have very similar 
leaflet arrangement, and have white indumentum on the lower leaflet surfaces, 
though in C. trachycoleus it is sparse and usually only present on young leaves. 
The leaf sheath with its abundant minute thorns forming a scabridity between the 
sparse large triangular thorns in C. trachycoleus and lacking such scabridity in 
C. caesius is the only reliable character for distinguishing the two species in the 
herbarium; yet in the field the two are seen to be remarkably distinct because of 
their differing habits. 


The striking differences in development of the suckering habit of these two 
rattans illustrates how important such differences may be to the silviculturalist; 
further studies of rattan suckering habits may produce other examples of closely 
related species differing in habit and are seen to be of great importance in a 
consideration of the ecology of rattans. 


ACKNOWLEDGEMENTS 


I should like to thank Sdr. Damhuri of the Herbarium Borgoriense for 
preparing the figures. 


REFERENCES 


BECCARI, O. (1913): Asiatic Palms — Lepidocaryeae, Part 1 The species of 
Calamus. Appendix. Ann. R. bot. Gard. Calcutta 11. 


HOLTTUM, R. E. (1955): Growth habit of monotyledons — variations on a 
theme. Phytomorphology 5 (4), 399-413. 


Synaptospory: a hypothesis 
A possible function of spore sculpture in pteridophytes 
by 


K. U. KRAMER 


Institute for Systematic Botany 
University of Ziirich 
Switzerland 


Summary 


Part of the spore output of some ferns is shed in groups of spores, sometimes still 
contained within the sporangium. It is theorized that spore sculpture, particularly a distinct, 
strongly sculptured perispore, plays a part in keeping, or bringing, spores together. This is 
important for increasing the chance for intergametophytic fertilization. The want of a 
pronounced spore sculpture in most epiphytic ferns and the strongly sculptured spores of 
heterosporous pteridophytes are brought in connexion with the phenomenon, tentatively 
called synaptospory. 


Until recently, pteridologists used to distinguish between fern taxa (genera 
or groups of higher rank) with a perispore and taxa without one, and much 
taxonomic value was attributed to this character (e.g... Kramer in Lanjouw & 
collab., 1968; Wagner, 1974). Through Lugardon’s fundamental work (e.g., 1971, 
1972a, 1974) it has now become known that the perispore is apparently of universal 
occurrence in ferns, and it has also been found in so-called fern allies (e.g., Lugardon 
1969, 1972b, 1973). It is therefore no longer correct to speak of ferns with and 
ferns without a perispore. Instead, one should distinguish between spores having a 
perispore that is firmly appressed to the exospore, and others where the perispore 
forms a distinct, usually strongly sculptured layer. To avoid this awkwardness in 
expression I shall continue to speak below of spores with and spores without a 
perispore, being, of course, fully aware of the incorrectness of these terms. 


In most cases a strongly developed surface sculpture of a spore was attributed 
to the perispore although in some cases the exospore was held responsible for the 
surface features. It now seems as if in all these cases a perispore is present, but 
where it seemed to be absent it firmly adheres to the exospore and cannot readily 
be distinguished except in cross-sections examined with the transmission electron 
microscope. ; 


Whatever the morphological interpretation, the function of this surface sculp- 
ture has remained a matter of conjecture, not to say an enigma, as it remains to this 
day, to a certain extent and largely as to details, in the pollen grains of seed plants. 
In the following I want to put forward a hypothesis that tries to account for the 
presence of spore sculpture in pteridophytes. I wish to stress that it may only be 
one of several possible functions, and that the attempt at an explanation is no 
more than a hypothesis, as for the moment it rests on too few observational data. I 
hope, though, that it may lead to further experimental work and will either be 
confirmed or, if disproved, will show to have been a stimulating error. 

When we review the “‘presence”’ or ‘‘absence” of a perispore (inverted commas 
because of the restrictions stated above). or, more generally, of a pronounced 
external sculpture in the spores of modern ferns, it becomes immediately apparent 
that large groups are without such sculpture and other, equally large and as a rule 


79 


80 Gardens’ Bulletin, Singapore — XXX (1977) 


very naturally defined ones, possess it. Wagner (l.c.) regarded the groups with a 
pronounced perispore as derived ones, and the perispore in general as a specializa- 
tion, whatever its function. It seems to me that there is another striking correlation, 
namely between the “‘presence” or “‘absence” of a perispore and the habitat of 
the fern, that has, to my knowledge, not been reported before. 


If we regard the spore wall of taxa of higher order among the modern ferns, 
it springs to the eye that epiphytic genera, genus groups, etc., in most cases have 
more or less smooth, ‘‘perisporeless’”” spores: Vittarioids, Davallioids, Grammi- 
tidoids, Polypodioids, Hymenophyllaceae. Minor exceptions can be found with ease, 
e.g. in the Polypodioids in Drymoglossum (see Wagner, l.c., fig. 2), in the 
Davallioids in Gymnogrammitis (see Ching, 1966, and Sen & Sen, 1971), etc., and 
there is one important major exception: the genus Elaphoglossum, This, though 
containing some terrestrial species, as do indeed most epiphytic groups, consists 
almost entirely of epiphytes; but its alliance is with a group of terrestrial or 
scandent genera known as the Lomariopsidoid ferns or Lomariopsidaceae*). 
‘*Perispore-less”’ spores occur, of course, in a considerable number of essentially or 
entirely terrestrial fern groups. Examples are Osmundaceae, Plagiogyriaceae and 
Dennstaedtioids. 


A second observation, of high relevancy in this context, was recently published 
by Schneller (1975). It has nearly always been assumed, explicitly or tacitly, that 
fern spores are scattered invidually as far as possible. Schneller showed that in 
Dryopteris spp. a not inconsiderable part of the spores is shed in groups of two, 
three, four, or more. 


According to Gastony’s findings (1974), in certain tree-ferns not the spore but 
the sporangium, with its complete spore-content, is the unit of dispersal. The spores 
germinate whilst held together by the sporangium. I have the strong impression 
that at least partial retention of spores in the sporangium. with subsequent shedding 
of the entire structure, is not a rare phenomenon in ferns. 


The pioneer work by Naf (1961) and Voeller (1971) has shown that the 
development of gametophytes and their sex organs is a process subject to physiologi- 
cal interaction between closely associated prothalli. These interactions greatly tend 
to favour intergametophytic fertilization and hence gene exchange. Various authors 
have reported on the much greater viability of fern sporophytes produced by 
intergametophytic as opposed to intragametophytic fertilization (e.g., Lloyd, 1974). 


Terrestrial ferns by and large may be supposed to grow in an essentially stable 
environment (there are, of course, exceptions like pioneers on bare soil, lava flows, 
etc.) where the rate of mortality of individual plants is relatively low. Once they 
are mature, individual plants often grow slowly and are supposed to be potentially 
very long-lived, e.g., in Osmunda regalis. Except when new habitats are colonized, 
sexually produced new generations will follow each other at rather long time 
intervals, and it must be of great importance for the plants to maintain their genetic 
variability, and to avoid the influence of recessive deleterious genes, by increasing 
the chance for cross-fertilization when new plants are sexually produced from 
prothalli. The coherence of some spores, whether through the action of the spore 
wall or by retention in the sporangium, must greatly enhance this chance, and 
external spore sculpture may be one mechanism by which the former is achieved. 
Spores from one sporangium will not be all genetically identical, unless the mother 
plant is one hundred percent homozygous. Schneller (1.c.) found many groups of 
more than four spores being dispersed as a unit, so the tetrad is certainly not the 


* On the evidence available at present I find myself unable to agree with Tutin (1964) and 
Pichi Sermolli (1968) who distinguish a family Elaphoglossaceae comprising only Elapho- 
glossum and satellite genera. 


Synaptos pory 81 


limit beyond which the spores do not cohere. And there seems no reason to suppose 
why spores produced by different plants from one colony, and even from different 
ones, could not meet after having been dispersed by air currents, runoff water, etc., 
for instance, in a surface-film of rainwater, cohere, and germinate together, with 
subsequent cross-fertilization. 


In contrast, most epiphytes inhabit a very unstable milieu, The branches and 
twigs they colonize have but a limited life-span, and they are thus bound to colonize 
new places continuously in order to survive as a species, Many do this by producing 
long-creeping, branching rhizomes, or sometimes runners, and such ferns very often 
inhabit the smaller branches and twigs of trees and shrubs, as I observed myself 
in some parts of tropical America, and as can be deduced from de la Sota’s diagram 
(1971, fig. 13) illustrating Costa Rican fern epiphytes; see also Johansson (1974). 
The epiphytes with short rhizomes, often more massive, or even forming so-called 
nests, favour the more stable habitats of the main trunks and crotches of larger 
branches. 


For colonizing new stations, these, as it were, pioneer epiphytes thus depend 
to a great extent on spore dispersal, which must be sufficient and efficient in order 
to ensure maintenance of a sufficiently large population, in spite of considerable 
turnover. This must largely be achieved by dispersing individual spores, and self- 
fertilization in lone prothalli originating from singly spread spores should be much 
more common in these plants. Indeed, coherence of spores in groups would impede 
dispersal. Wagner (l.c.) suggested that the lack of a perispore in Davallioid ferns 
should be regarded as a result of secondary loss of this organ, and if that is true, 
it might be regarded as an adaptation to epiphytism. With the comparatively rapid 
succession of sexually produced generations, occasional cross-fertilization would 
suffice for maintaining genetic variability. 


In this connexion it is, I think, significant that so very few natural hybrids of 
epiphytic fern species have been reported, in contrast to the many terrestrial hybrids 
known. This may be due in part to the physical inability of the researcher to 
sample populations of epiphytes in the same way as of terrestrial or epilithic 
species, but it can hardly account for the fact as a whole. The regular development 
of single, self-fertilized prothalli may well be the reason for a greatly reduced 
chance for interspecific cross-fertilization. 


The phenomenon of coherence of spores by means of cohering or interlocking 
surface structures may be called “‘synaptospory’’, analogous to what has been called 
“synaptospermy”’ in seed plants, after Murbeck (see, for instance, Zohary, 1962, 
p. 180, and van der Pijl, 1969, p. 77). Further research will have to show whether it 
exists as a widespread phenomenon. 


One general feature observed in terrestrial heterosporous lycopodiophytes 
seems to point in the same direction. The great majority of Selaginella species, and 
many species of the partly terrestrial genus Jsoetes, have strongly sculptured spores. 
For illustrations, see, for instance, Knox (1950) and Hellwig (1969). For sexual 
reproduction these plants are, of course, entirely dependent on germination of 
spores producing gametes of opposite sex in very close proximity. The surface 
sculpture may here, too, serve to enhance the chance for this to occur. 


Equisetum is probably another case in point. The perispore here dissolves into 
hygroscopically moving bands, somewhat unfortunately termed elaters, which loosen 
the spore mass but also become entangled, leading to distribution of spores in groups. 
The prothalli are at least very often functionally unisexual (Sporne, 1952, p. 112; 
Duckett & Duckett, 1974). 


I am indebted to Mr. E. Hennipman, Leiden, for stimulating discussions. 


82 Gardens’ Bulletin, Singapore — XXX (1977) 


References 


Ching, R. C. 1966. Gymnogrammitidaceae Ching, a new fern family. Acta Phytotax. 
Sin. 11: 11-16. 


Duckett, J. G., & Duckett, A. R. 1974. The ecology of Equisetum gametophytes. 
Am. J. Bot. 61: 36 [brief report). 


Gastony, G. J. 1974. Spore morphology in the Cyatheaceae, I. The perine and 
sporangial capacity: general considerations. Am. J. Bot, 61: 672-680. 


Hellwig, R. L. 1969. Spores of the heterophyllous Selaginellae of Mexico and 
Central America. Ann. Missouri Bot. G. 56: 444-464. 


Johansson, D. 1974. Ecology of vascular epiphytes in West African rain forest. 
Acta Phytogeogr. Suec. 59: 1-129. 


Knox, E, M. 1950. The spores of Lycopodium, Phylloglossum, Selaginella and 
Isoetes, and their value in the study of microfossils of Paleozoic age. Trans. 
Bot. Soc. Edinb. 35: 211-357; 12 plates. 


Kramer, K. U. 1968. Pteridophyta, in: J. Lanjouw & collab. Compendium van de 
Pteridophyta en Spermatophyta. Oosthoek, Utrecht. 342 pp. 


Lloyd, R. M. 1974, Genetic and mating system studies in ferns: systematic and 
evolutionary implications, Internat. Organiz. Plant Biosyst. Newsletter 9: 2-14. 


Lugardon, B. 1969. Sur la structure fine des parois sporales d’Equisetum maximum 
Lamk. Pollen et Spores 11: 449-474. 


1971. Contribution a la connaissance de la morphogénése et de la 
structure des parois sporales chez les Filicinées isosporées. Thése, Université 
de Toulouse. 257 pp., 51 plates. 


1972a. La structure fine de l’exospore et de la périspore des 
Filicinées isosporées, I — Généralités. Eusporangiées et Osmundales. Pollen 
et Spores 14: 227-261. 


1972b. Sur la structure fine et la nomenclature des parois micro- 
sporales chez Selaginella denticulata (L.) Link et S. selaginoides (L.) Link. 
C. R. Acad. Sci. Paris Sér. D 274: 1656-1659. 


1973, Nomenclature et structure fine des parois acétorésistantes des 
microspores d’Isoetes. C. R. Acad. Sci. Paris Sér. D 276: 3017-3020. 


1974, La structure fine de l’exospore et de la périspore des Filicinées 
isosporées. II. Filicales. Commentaires. Pollen et Spores 16: 161-226. 


Naf, U. 1961. Mode of action of an antheridium-inducing substance in ferns. Nature 
189: 900-903. 


Pichi Sermolli, R. E. G. 1968. Adumbratio Florae Aethiopicae 15. Elaphoglossaceae. 
Webbia 23: 209-246. 


van der Pijl, L. 1969. Principles of dispersal in higher plants. Springer-Verlag, 
Berlin. 154 pp. 


Schneller, J. 1975. Untersuchungen an einheimischen Farnen, insbesondere der 
Dryopteris filix-mas-Gruppe. 3. Teil. Oekologische Untersuchungen. Ber. 
Schweiz. Bot. Ges. 85: 110-159. 


Sen, T., & Sen, U. 1971. Morphology and anatomy of the fern genus Gymnogram- 
mitis. Ann. Bot. 35: 229-235. 


Synaptospory 83 

de la Sota, E. R. 1971. El epifitismo y las pteriddfitas en Costa Rica (América 
Central). Nova Hedwigia 21: 401-465. 

Sporne, K. R. 1962. The morphology of pteridophytes. Hutchinson, London. 192 pp. 


Tutin, T. G. 1964. Elaphoglossaceae, in: T. G. Tutin, V. H. Heywood & collab., 
Flora Europaea Vol, 1. Cambridge. xxxii + 464 pp. 


Voeller, B. 1971. Developmental physiology of fern gametophytes: relevance for 
biology. BioScience 21: 266-270. 


Wagner, W. H. Jr. 1974. Structure of spores in relation to fern phylogeny. Ann. 
Missouri Bot. G. 61: 332-353. 


Zohary, M. 1962. Plant life of Palestine. Ronald Press, New York. 262 pp. 


Vol. 20008 vojst; fatale nisi 


“ih neaolvda (risi qi noiralin ni enone. oe BU 
. 6h: 


ca, SA® abo ¥ wet. ever blend per her 15 wi Da 


a: 


WM. 


» *)* " 
©, 


Use fie (Mie iY ies RSS 


pats a 

a » ~ 

a) ‘os ~ 0 ae a S Mae on, 
oa Nees eae. ee : 


~ 


Spore Morphology of Malayan Dennstaedtiaceae sensu Holttum 
by 
E. SoEPADMO & E. E. KHoo 


Department of Botany 
University of Malaya 
Kuala Lumpur 


Abstract 


Spore morphology of 28 species belonging to 28 genera included by Holttum (1954) in 
the Dennstaedtiaceae are described. Amongst the Malaysian taxa there are at least seven 
distinct spore-types. The disposition of the Malaysian genera in these spore-types does not 
conform with any of the existing systems of classification. Only spores of those genera 
assigned by Holttum in the subfamilies Davallioideae, Dryopteridoideae and Tectarioideae 
show some degree of uniformity. 


Introduction 


The Dennstaedtiaceae as originally defined by Holttum (1947, 1949, 1954) 
is a very large family of modern ferns, containing very diverse groups of genera 
and species. In the Peninsular Malaysia alone the family comprises not less than 
48 genera and 203 species out of the total 124 genera and 389 species of its fern- 
flora. As has been shown in the excellent review by Pichi Sermolli (1973) and 
Mickel (1973), there seems to be a great controversy over the delimitation and 
classification of the taxa involved. Many authors, such as Christensen (1938), 
Ching (1940), Copeland (1947), Reimers (1954), Pichi Sermolli (1958), Nayar 
(1970) and others placed the 48 Malaysian genera involved in 5 or 10 separate 
families, whereas Holttum (1954) included these genera in 11 subfamilies of the 
Dennstaedtiaceae, though later (1973) he more or less agrees with Pichi Sermolli’s 
system of classification. Furthermore, it is also evident that there is a great 
difference of opinion with regard to the disposition and affinity of the genera 
within each family or subfamily recognised by the above authors. 


In an attempt to introduce a more natural system of classification of these 
taxa, many pteridologists (e.g. Atkinson, 1973; Lovis, 1973; Manton, 1961; Mickel, 
1973; Sen & Sen, 1973; Sledge, 1973; Swain & Cooper-Driver, 1973; Van Cotthem, 
1973; Walker, 1973, and others) have used modern and advanced techniques in 
their studies, previously unexploited by those who are responsible in drawing up 
the existing schemes of classification. Amongst these modern techniques are: 
anatomical, cytological, phytochemical, biochemical, developmental and palyno- 
logical. 


Of the palynological aspects, Erdtman & Sorsa (1971) have admirably 
summarised our present knowledge on the spore morphology of those genera 
included in the Dennstaedtiaceae by Holttum. This work shows that of the total 
3052 species so far recognised in the 48 genera occurring in Malaysia (Willis, 
1973), spore descriptions of only 449 (= 14.7%) have been published by Harris 
(1955), Erdtman (1957), Nayar & Devi (1963, 1964a, 1964b, 1966, 1967, 
1968a, 1968b), Nayar & Kaur (1963), Tardieu-Blot (1963a & b, 1965), and 
several others. Of these descriptions only a few were based on Malaysian specimens. 


85 


86 Gardens’ Bulletin, Singapore — XXX (1977) 

The present paper describes the spores of 28 species belonging to 28 genera, 
and is intended as a starting point for more intensive works to be done in the 
future. | 


Species Investigated 


1. DENNSTAEDTIOIDEAE 6. ASPLENIOIDEAE 
Hypolepis bivalvis v.A.v.R. Asplenium nidus L. var. musifolia 
Microlepia speluncae (L.). Moore (Sm.) Bedd. 
Orthiopteris kingii (Bedd.) Holtt. gv igi comes 
2. LINDSAEOIDEAE Blechnum orientale L. 
Isoloma divergens (Hook. f. & Grev.) Brainea insignis (Hook. f.) Sm. 
Sm. 


8. LOMARIOPSIDOIDEAE 
Bolbitis heteroclita (Pr.) Ching 
Egenolfia appendiculata (Willd.) Sm. 
Elaphoglossum callifolium (Bl.) Moore 


Lindsaea scandens Hook. f. 
Sphenomeris chusana (L.) Copel. 
Tapeinidium pinnatum (Cav.) Christ. 


3. DAVALLIOIDEAE Lomagramma sumatrana v.A.V.R. 
Davallia divaricata Bl. Teratophyllum aculeatum (Bl.) Mett. 
Humata heterophylla (Sm.) Desv. var. montanum Holtt. 

4. OLEANDROIDEAE 9. DRYOPTERIDOIDEAE 
Nephrolepis biserrata (Sw.) Schott. Dryopteris sparsa (Don) O. Ktze. 
Oleandra pistillaris (Sw.) Christ. 10. TECTARIOIDEAE 

5. PTERIDIOIDEAE Arcypteris irregularis (Pr.) Holtt. 
Acrostichum aureum L. Heterogonium pinnatum (Copel.) 
Histiopteris stipulacea (Hook. f.) Holtt. 

Copel. 11. ATHYRIOIDEAE 


Pieris vittata L. 


Athyrium pinnatum (Blanco) Copel., 
Stenochlaena palustris (Burm.) Bedd. 


Cystopteris tenuisecta (Bl.) Mett. 
Materials and Methods 


In this study both fresh and herbarium materials were used. The fresh 
specimens were collected from various localities in Selangor and Pahang, and were 
identified with the help of Holttum’s keys (1954). Voucher specimens are now 
deposited in the Herbarium, Department of Botany. University of Malaya, Kuala 
Lumpur (KLU). Dried and preserved materials were obtained from herbarium 
specimens available in the above Herbarium. 


Spores containing materials were acetolysed, stained lightly with safranin, and 
mounted in glycerin jelly. The slides were then sealed permanently with paraffin 
wax. In some species, during acetolysis the perine was easily dissolved and the 
spores become shrivelled. In such cases, a lesser amount of concentrated sulphuric 
acid was added in the acetolysing mixture and heating was skipped. Although by 
this treatment, the spores did not become transparent, the perine remained intact 
and could be studied. Observation, measurement and photomicrography was 
carried out with a Leitz Dialux Microscope fitted with occular micrometer, 
Combiphot Automatic Camera System, achromatic condensor with N.A, 0.90, 
phaco achromatic objectives 40x /0.65 and 100X /1.30, and halogen lamp-housing 
model 100 Z. 


For each species, measurement is based on 50 readings. This measurement is 
expressed in the text as length of polar axis (P) x longitudinal equatorial diameter 
(El) X transverse equatorial diameter (E2) in monolete-bilateral spores and as 
P x E1l in trilete-tetrahédral spores. All measurements are exclusive of perine and 
any other processes. : 


Malayan Dennstaedtiaceae | 87 


Description of Spores 


DENNSTAEDTIOIDEAE 


1. Hypolepis bivalvis (Gunong Brinchang, Perak; Poore KLU 468; Plate 8, 

Figs. 4-7). 

Spores monolete-bilateral, 30-46 x 48-61 x 33-46 ». Amb oblong; proximal 
face flat, distal face hemispherical. Laesura 36-40 yp, margins thickened into 
protruding lips about 4 to 7 » broad on either side of the laesura. Exine 
2 p» thick, sexine thicker than nexine, with fine spinulose pattern. Perine thin and 
provided with minute spine-like processes. 


2. Orthiopteris kingii (Cameron Highlands, Pahang; Turnau KLU 2936; Plate 2, 

Fig 8). 

Spores trilete-tetrahedral, 29-32 x 31-39 ». Amb triangular in outline, with 
convex sides and broadly rounded corners. Proximal face conical, distal face 
convex to hemispherical. Laesura 15 yw, margins thickened into lips about 
3 » broad and tapering towards the tips. Exine 2 » thick, sexine nearly as thick 
as nexine, smooth. Perine absent. 


3. Méicrolepia speluncae (Templer Park, Selangor; Abd. Samat b, Abdullah KLU 
2686; Plate 2, Figs. 6-7). 


Spores trilete-tetrahedral, 25-33 x 30-33 yw. Amb triangular with strongly 
concave sides and broadly rounded corners, Proximal face slightly convex to 
flat; distal face hemispherical. Laesura 13 y», margin thickened into lips of 
2 » broad and tapering towards the tips. Exine 2 » thick, sexine as thick as nexine, 
densely granulose. Perine absent. 


LINDSAEOIDEAE 


4. Lindsaea scandens (Gunong Ulu Kali, Pahang; Abd. Samat b. Abdullah KLU 

1846; Plate 2, Figs. 2-3). 

Spores trilete-tetrahedral, 22-25 x 24-27 uu. Amb triangular, with slightly 
convex to straight sides. Proximal face conical with angular pole; distal face 
convex. Laesura 13 », margins thickened into lips of 1 yw broad. Exine 
1 uw thick, not clearly subdivided into sexine and nexine, densely granulose. Perine 
absent, 


5. Sphenomeris chusana (Fraser’s Hills, Pahang; Turnau KLU 2701; Plate 9, 

Fig. 3). 

Spores monolete-bilateral, 43-63 x 65-76 x 40-51 u. Amb oblong. Proximal 
face flat; distal face hemispherical. Laesura 51 u, margins thickened into protruding 
lips of 8 » broad. Exine 2 » thick, sexine much thicker than nexine, smooth. 
Perine smooth and thin, partially peeling off on acetolysis. 


6. Isoloma divergens (Genting Simpah, Selangor; Mohd. Kassim b. Rajab KLU 
139; Plate 2, Figs. 4-5). 


Spores trilete-tetrahedral, 19-22 x 20-25 uw. Amb triangular with straight to 
slightly convex sides and broadly rounded corners. Proximal face conical with 
angular pole; distal face convex. Laesura 12 », margins thickened into lips of 
1 » broad. Exine 1.5 » thick, sexine much thicker than nexine, smooth. Perine-like 
layer present, smooth, very closely adherent to the exine, and disintegrates very 
easily on acetolysis. 


7. Tapeinidium pinnatum (Fraser’s Hills, Pahang; Khoo Eng Ee KLU 12307; 
Plate 9, Fig. 4). 


88 Gardens’ Bulletin, Singapore — XXX (1977) 


Spores monolete-bilateral, 26-32 x 28-46 x 27-32 ». Amb. oblong. Proximal 
face flat to slightly convex; distal face hemispherical. Laesura 32 y», margins 
thickened into protruding lips of 6 » broad. Exine 1.5 » thick, sexine thicker 
than nexine, smooth. Perine thin with fine reticulate pattern, disintegrates easily 
on acetolysis. 


DAVALLIOIDEAE 


8. Davallia divaricata (Fraser’s Hills, Pahang; Khoo Eng Ee KLU 12309; Plate 

6, Figs. 1-3). 

Spores monolete-bilateral, 27-35 x 36-50 x 24-34 ». Amb oblong. Proximal 
face concave; distal face convex. Laesura 28 », margins slightly thickened into lips 
of 2 » broad. Exine 2.5 » thick, sexine thicker than nexine, verrucose to rounded 
tuberculose; verrucae or tubercles crowded, 5-7 p» in diameter and 2 uw» high. 
Perine absent. 


9. Humata heterophylla (Lombong, Johore; Abd. Samat b. Abdullah KLU 1849; 
Plate 6, Figs. 4-8). 


Spores monolete-bilateral, 22-36 <x 38-43 x 28-32 ». Amb oblong. Proximal 
face flat; distal face hemispherical. Laesura 18 », margins slightly thickened into 
lips of 1 » broad. Exine 3 » thick, sexine thicker than nexine, verrucose-tuber- 
culose; tubercles crowded, round, 2 » high, 5-7 » broad and 10-16 wp long. 
Perine absent. 


OLEANDROIDEAE 


10. Oleandra pistillaris (Fraser’s Hills, Pahang; Khoo Eng Ee KLU 12310; 

Plate 4, Figs. 5-6). 

Spores monolete-bilateral, 19-26 x 27-32 x 17-23 ». Amb oblong. Proximal 
face flat; distal face hemispherical. Laesura 17 », margins slightly thickened into 
lips of 1 » broad. Exine 4-5 yp» thick, sexine thicker than nexine, smooth. Perine 
loosely wrinkled, anastomosing to form a reticulate patterns; folds about 4 » high. 
spinulose; spinules minutes, less than 1 p» high, with pointed tips. Perine disinte- 
grates very easily on acetolysis. 


11. Nephrolepis biserrata (Damansara Road, Kuala Lumpur, Selangor; Khoo Eng 

Ee KLU 12305; Plate 7, Figs. 3-4). 

Spores monolete-bilateral, 25-30 x 33-40 x 23-29 ». Amb oblong. Proximal 
face flat to slightly concave; distal face convex. Laesura 20 u, margins thickened 
into lips of 1 » broad, tapering towards the tips. Exine 2 » thick, sexine thicker 
than nexine, verrucose to tuberculose; verrucae 1 p» high and 46 y» broad. 
Perine thin, more or less verrucose and easily disintegrates on acetolysis. 


PTERIDIOIDEAE 


12. Histiopteris stipulacea (Fraser’s Hills, Pahang; Khoo Eng Ee KLU 12306; 

Plate 7, Figs. 1-2). 

Spores monolete-bilateral, 38-48 x 59-65 x 43-46 ». Amb oblong. Proximal 
face slightly concave; distal face convex. Laesura 40 p, margins thickened slightly 
into lips of 6-10 » broad and tapering towards the tips. Exine 5 p» thick, sexine 
thicker than nexine, densely verrucose-tuberculose; verrucae or tubercles granulose, 
5 p» high, 6 » broad, and 8-22 » long. Perine absent. 


13. Pteris vittata (Fraser’s Hills, Pahang; Khoo Eng Ee KLU 12306; Plate 1, 
Figs. 1-3). 


Spores trilete-tetrahedral, 48-53 x 61-65 yw. Amb triangular with nearly 
straight sides and rounded corners. Proximal face flat to slightly convex; distal 


Malayan Dennstaedtiaceae 89 


face hemispherical. Laesura 27 », margins thickened into lips of 3 « broad and 
tapering towards the tips. Exine 3 « thick, sexine much thicker than nexine, densely 
rugulose; rugulae up to 6 u tall, sometimes coalescing to form a coarse reticulum. 
On the proximal face there are 3 ridges parallel and close to the laesura arms. 
Equator collar present; this is a collar-like ridge which protrudes as a flange 
around the spores, separating the proximal and the distal parts of the spores, up 
to 7 » high, often interrupted. Perine absent. 


14. Acrostichum aureum (Lukut, Negeri Sembilan; Turnau KLU 2925: Plate 2, 

Fig. 1). 

Spores trilete-tetrahedral, 42-49 x 50-60 «. Amb rounded triangular with 
slightly convex sides and broadly rounded corners. Proximal face conical with 
straight sides and angular pole; distal face hemispherical. Laesura 21 «, margins 
thickened into lips of 1 « broad and tapering towards the tips. Exine 2 « thick, 
sexine thicker than nexine, granulose. Perine not seen. 


15. Stenochlaena palustris (Damansara Road, Kuala Lumpur, Selangor; Khoo 
Eng Ee KLU 12311; Plate 7, Figs. 5-6). 


Spores monolete-bilateral, 30-37 x 39-53 xX 30-36 x. Amb oblong. Proximal 
face flat to slightly concave; distal face hemispherical. Laesura 29 xu, margins 
thickened into lips of 1 « broad. Exine 3 uw thick, sexine thicker than nexine, 
provided with conical tubercles of 2 » tall with rounded tips and base diameter 
of 2 uw. Perine absent. 


ASPLENIOIDEAE 


16. Asplenium nidus var. musifolia (Damansara Road, Kuala Lumpur; Khoo Eng 
Ee KLU 12304; Plate 3, Figs. 1-2). 


Spores monolete-bilateral, 27-36 x 30-SO X 27-32 u. Amb oblong. Proximal 
face flat to slightly concave; distal face hemispherica]. Laesura 20 xu, margins 
thickened slightly into lips of 3 u broad. Exine 1 w thick, sexine as thick as 
nexine, smooth. Perine finely granulose, wrinkled into discontinous, closely sinuous 
folds protruding up to 3 » from the exine surface, with smooth crests. 


BLECHNOIDEAE 


17. Blechnum orientale (Ulu Gombak Road, Selangor; Khoo Eng Ee KLU 
12308; Plate 3, Figs. 3-4). 


Spores monolete-bilateral, 34-39 x 44-51 X 3646 u. Amb oblong. Proximal 
face flat; distal face hemispherical. Laesura 29 », margins thickened into lips 
3 » broad and tapering towards the tips. Exine 1 « thick, sexine as thick as 
nexine, smooth. Perine sparsely granulose, loosely folded into sparse, thin indistinct 
folds protruding up to 3 u« from the exine surface, with smooth crests: disintegrates 
easily on acetolysis. 


18. Brainea insignis (Tanjong Selantai, Johore; Abd. Samat b. Abdullah KLU 
1869; Plate 10, Figs. 4-6). 


Spores monolete-bilateral, 31-42 x 35-SO X 29 43 x. Amb oblong to sub- 
spherical. Proximal face slightly convex; distal face hemispherical. Laesura 27 x, 
margins thickened into lips of 1 « thick and tapering towards the tips. Exine 
2 m thick, sexine as thick as nexine, smooth. Perine densely granulose, closely 
adherent to the exine nearly without folds or only with one or two indistinct short 
folds which protrude up to 2 « from the exine surface; disintegrates easily on 
acetolysis. 


90 Gardens’ Bulletin, Singapore — XXX (1977) 


LOMARIOPSIDOIDEAE 


19. Egenolfia appendiculata (Kuah, Langkawi, Kedah; Abd. Samat b. Abdullah 

KLU 2316; Plate 10, Fig. 3). 

Spores monolete-bilateral, 32-36 x 31-39 x 26-37 ». Amb subspherical to 
circular in outline. Proximal face flat to slightly convex; distal face hemispherical. 
Laesura 19 yp, slightly thickened into lips of 1 » broad; in some spores forked in 
one end. Exine 1 » thick, sexine as thick as nexine, smooth. Perine with finely 
reticulate patterns, loose, lightly folded into very many elongated sharp ridges most 
of which radiate from the centre to the periphery of the spores and protruding up 
to 7 » from the exine surface; disintegrates on acetolysis. 


20. Bolbitis heteroclita (Jengka Forest Reserve, Pahang; Turnau KLU 2894; 

Plate 10, Figs. 1-2). 

Spores monolete-bilateral, 31-37 x 35-42 x 35-38 yw. Amb oblong to 
subspherical. Proximal face flat to convex: distal face hemispherical. Laesura 21 uy, 
margins slightly thickened into smooth lips of 1 » broad. Exine 1 yw» thick, sexine 
as thick as nexine, smooth. Perine sparcely covered with minute spinules, highly 
folded into sinuous and elongated ridges radiating from the centre to the periphery 
of the spores; folds protrude up to 12 u from the exine surface, with undulating 
crests; disintegrates on acetolysis. 


21. Teratophyllum aculeatum var. montanum (Tringkap, Pahang Abd. Samat b. 
Abdullah KLU 2288; Plate 5, Figs. 1-3). 


Spores monolete-bilateral, 47-60 Xx 66-71 uw (polar view not obtained). 
Proximal face convex to hemispherical; distal face hemispherical. Laesura 27-30 up. 
Exine very thin, less than 1 » thick, not subdivided into distinct sexine and nexine. 
Perine densely crowded with minute spinules less than 1 » high, very sparsely 
folded into faint folds which protrude up to 19 » from the exine surface; with 
smooth crests; cracked up on acetolysis, 


22. Lomagramma sumatrana (22nd mile Bentong Road, Selangor; Turnau KLU 
2914; Plate 9, Figs. 1-2). 


Spores monolete-bilateral, 31-43 x 50-58 x 35-40 ». Amb oblong. Proximal 
face flat to slightly concave; distal face hemispherical. Laesura 29 ,, margins 
slightly thickened into lips of 1 » broad. Exine 2 » thick, sexine slightly thinner 
than nexine, finely granulose to spinulose. Perine absent. 


23. Elaphoglossum callifolium (Tanah Rata, Pahang; Mahmud b. Sider KLU 

4612; Plate 4, Figs. 1-2). 

Spores monolete-bilateral, 18-30 x 23-40 x 18-22 ». Amb oblong. Proximal 
face flat to slightly concave; distal face hemispherical. Laesura 18 yw. Exine 
1 » thick, sexine and nexine are not clearly differentiated, smooth. Perine minutely 
spinulose, sparsely folded into thin, sharp sinuous ridges protruding up to 7 » from 
the exine surface; disintegrates on acetolysis. 


DRYOPTERIDOIDEAE 
24. Dryopteris sparsa (Maxwell Hills, Perak; Evans KLU 2493; Plate 3, 
Figs. 5-6). . 


Spores monolete-bilateral, 22-27 x 38-46 x 28-31 ». Amb oblong. Proximal 
face flat to slightly concave; distal face convex. Laesura 20 », margins thickened 
very slightly into lips of 1 » broad. Exine 1 ,» thick, sexine as thick as nexine, 
smooth. Perine smooth, fairly closely adherent to the exine, folded into lobate 
crowded ridges protruding up to 11 » from the exine surface; with many undulating 
crests; disintegrates on acetolysis. 


Malayan Dennstaedtiaceae 9] 


TECTARIOIDEAE 


25. Arcypteris irregularis (Ulu Gombak Road, Selangor; Wee & Lim KLU 1975; 
Plate 5, Figs. 4-6). 


Spores monolete-bilateral, 20-29 x 32-39 x 23-26 ». Amb oblong. Proximal 
face flat; distal face hemispherical. Laesura 20 », margins faintly thickened into 
lips of 1 » broad. Exine 1.5 » thick, sexine as thick as nexine, smooth. Perine with 
very minute spinules, folded into a few elongated sharp, thin ridges radiating from 
the centre to the periphery of the spores; folds protrude up to 10 » from the exine 
surface; disintegrates easily on acetolysis, 


26. Heterogonium pinnatum (Templer Park, Selangor; Abd. Samat b. Abdullah 
KLU 2636; Plate 4, Figs. 3-4). 


Spores monolete-bilateral, 283-40 x 40-50 x 33-36 u. Amb oblong. Proximal 
face flat to slightly concave; distal face hemispherical. Laesura 27 uw, margins 
faintly thickened into lips of 1 » broad. Exine 2 wu thick, sexine as thick as 
nexine, smooth, breaking up on acetolysis. Perine minutely spinulose (spines less 
than 1 » tall), highly folded up into elongated, closely sinuous ridges protruding 
up to 6 » from the exine surface; disintegrates on acetolysis. 


ATHYRIOIDEAE 
27. Athyrium pinnatum (Genting Simpah, Selangor; Turnau KLU 2785; Plate 
9, Figs. 5-6). 


Spores monolete-bilateral, 20-27 x 33-44 x 24-25 uw. Amb oblong. Proximal 
face flat to convex; distal face hemispherical. Laesura 23 yw, margins faintly 
thickened into lips of 1 » broad. Exine 2 yw thick, sexine as thick as nexine, 
smooth. Perine faintly granulose, closely adherent to the exine, with a few folds; 
folds appear as irregular ridges protruding up to 2 u» from the exine surface; 
disintegrates on acetolysis. 


28. Cystopteris tenuisecta (Cameron Highlands, Pahang: Poore KLU 256; Plate 

8, Figs 1-3). 

Spores monolete-bilateral, 30-45 x 45-60 x 31-45 u. Amb oblong. Proximal 
face flat to slightly concave; distal face hemispherical. Laesura 30 yu, margins 
slightly thickened into lips of 1 « broad. Exine 2 ,» thick, sexine thicker than 
nexine, densely finely spinulose. Perine folded; folds anastomose to form dense 
reticulum, provided with numerous rod-like protuberances about 3-4 .» high; tips 
of rods peltately lobed. 


Discussion 


Spore morphology of 28 Malaysian species included by Holttum (1954) in 
the Dennstaedtiaceae shows a great diversity of form, size and structure. Based 
on whether the spores are monolete-bilateral or trilete-tetrahedral, with or without 
perine, and perine if present smooth or variously patterned, etc., there seem to be 
at least seven distinct types of spores. These spore-types are: 


Type-1: Spore trilete-tetrahedral, provided with well-developed equatorial collar 
(Pteris vittata). 


Type-2: Spore trilete-tetrahedral, without equatorial collar, without or occasionally 
with a thin spinulose perine (Acrostichum aureum, Isoloma divergens, 
Lindsaea scandens, Microlepia speluncae, and Orthiopteris kingii). 


Type-3: Spore monolete-bilateral, provided with folded or reticulate perine, plano- 
convex or concave-convex in equatorial view (Asplinium nidus var. 
musifolia, Blechnum orientale, Dryopteris sparsa, Elaphoglossum calli- 
folium, Heterogonium pinnatum, Oleandra pistillaris, Arcypteris irregu- 
laris, and Teratophyllum aculeatum). 


92 Gardens’ Bulletin, Singapore — XXX (1977) 


Type-4: Spore monolete-bilateral, without or occasionally with very thin perine, 
exine provided with low, flattish, roundish or conical protuberances 
(Davallia divaricata, Histiopteris stipulacea, Humata heterophylla, 
Nephrolepis biserrata, and Stenochlaena palustris). 


Type-5: Spore monolete-bilateral, plano-convex, perine provided with spine-like 
or rod-like blunt processes (Cystopteris tenuisecta, Hypolepis bivalvis). 


Type-6: Spore monolete-bilateral, concave-convex or plano-convex, with or with- 
out perine; perine if present thin, hardly folded and easily dissolved on 
acetolysis (Athyrium pinnatum, Lomagramma sumatrana, Sphenomeris 
chusana, and Tapeinidium pinnatum). 


Type-7: Spore monolete-bilateral, biconvex or sometimes plano-convex, provided 
with folded, reticulate or finely patterned perine (Bolbitis heteroclita, 
Brainea insignis, and Egenolfia appendiculata). 


In the Dennstaedtioideae, Holttum included Dennstaedtia, Hypolepis, Micro- 
lepia and. Orthiopteris. Hypolepis with a monolete-bilateral spore of type-5 seems 
to be out of place in this subfamily, since the other three genera have trilete- 
tetrahedral spores of type-2. Furthermore, the perine in Hypolepis is provided with 
spine-like or rod-like minute processes, while the perine of Dennstaedtia and 
Orthiopteris, if present, is not so patterned, and in Microlepia the spores are with- 
out perine. Erdtman & Sorsa (1971) included these four genera in the family 
Pteridaceae in which 52 out of 65 genera have exclusively trilete-tetrahedral 
spores, 7 possess mainly trilete but occasionally also monolete spores, and 6 
genera with exclusively monolete-bilateral spores. Apart from Hypolepis, the other 
five genera which have exclusively monolete-bilateral spores are:— Humblotiella, 
Tapeinidium, Sphenomeris, Paesia, and Lonchitis. Perine of these genera, if present, 
is very thin, hardly folded and is easily detached from the exine on acetolysis, 
unlike that of Hypolepis. In Holttum’s classification, Tapeinidium and Sphenomeris, 
on the other hand, were grouped together with Lindsaea, Schizoloma and Isoloma 
in the Lindsaeoideae. Spore morphology does not support this grouping, since 
both Isoloma and Lindsaea have a trilete-tetrahedral spore of type-2, whereas 
Sphenomeris and Tapeinidium have a monolete-bilateral spore of type-6. 


Of the Davallioideae, the four genera (i.e. Araiostegia, Davallia, Humata, 
and Leucostegia) assigned by Holttum (1954) in this subfamily have a more or 
less similar spore-type, namely monolete-bilateral, without perine, and the exine 
is provided with low flattish verrucae or tubercles (spore type-4). For these 
features, the spores of Nephrolepis, Histiopteris and Stenochlaena agree very well. 
Holttum (1954), on the other hand, placed Nephrolepis together with Oleandra in 
the Oleandroideae. Spores of Oleandra are, however, different from those of 
Nephrolepis by being provided with folded perine, the detailed structure of which 
matches very well with that of Elaphoglossum, Arcypteris, and Teratophyllum (spore 
type-3). Histiopteris and Stenochlaena together with Acrostichum, Pteridium and 
Pteris were included by Holttum (l.c.) in the Pteridioideae. Acrostichum and 
Pteridium possess a trilete-tetrahedral spores of type-2, whereas Pteris has spores 
of type-1. 

The genus Asplenium (included by Holttum in the Asplenioideae) has spores 
which are very much similar to those of Blechnum and Heterogonium (spore 
type-3). However, Blechnum was assigned by Holttum in the Blechnoideae together 
with Brainea and Woodwardia. While spores of Woodwardia match very well with 
those of Blechnum, that of Brainea being monolete-bilateral, biconvex and provided 
with folded, granulose-reticulose perine, agree well with the spores of Bolbitis and 
Egenolfia (spore type-7). The last two mentioned genera were placed together 
with Elaphoglossum, Lomagramma, Lomariopsis and Teratophyllum in the 
Lomariopsidoideae by Holttum. While spores of Elaphoglossum and Teratophyllum 


Malayan Dennstaedtiaceae 93 


are very much similar to that of Oleandra and Tectaria (spore type-3), those of 
Lomagramma, being without perine, agree well with the spores of Athyrium, 
Sphenomeris and Tapeinidium (spore type-6). According to Erdtman & Sorsa 
(1971), some species of Lomariopsis possess perinous spores similar to 
those described as type-3, while others have a spore of type-6. 


In the Dryopteridoideae, Holttum assigned Acrophorus, Diacalpe, Didymo- 
chlaena, Dryopteris, Polystichum and Polystichopsis. These genera have more or 
less uniform spores which match very well with those of Asplenium, Blechnum 
and Heterogonium (Spore type-3). Similarly, the spores of ten genera included 
by Holttum in the Tectarioideae (Arcypteris, Ctenitis, Cyclopeltis, Heterogonium, 
Hypodematium, Lastreopsis, Pleocnemia, Pteridrys, Quercifilix and Tectaria) 
agree well with each other and more or less similar to those of spore type-3. 

Of the Athyroideae, Holttum place two Malaysian genera, namely Athyrium 
and Cystopteris. As has been indicated above, the spores of Athyrium belong to 
spore type-3, whereas those of Cystopteris to spore type-5. 

The above discussion seems to indicate that apart from the Davallioideae, 
Dryopteridoideae and Tectarioideae, the other subfamilies of the Dennstaedtiaceae 
as defined by Holttum, show very diverse types of spores and that spore morpho- 
logy does not lend any support to the disposition of various genera either in the 
different subfamilies as recognised by Holttum (1954) or in the various families as 
subscribed by Copeland (1947), Reimers (1954), Pichi Sermolli (1958) and 
others. 

However, this does not imply that the classification of the genera included by 
Holttum in the Dennstaedtiaceae should match well with the spore types, but 
rather that it demonstrates that more palynological studies, especially on those of 
the tropical taxa should be done, so that morphological characters of the spores 
could be considered along with other attributes in constructing a more natural 
system of classification. 

Finally, it should be emphasised here that, since the number of species of 
which the spores have been studied in detail, is very small, and that, it is also well 
known that even within a single genus the spore morphology may vary conside- 
rably, the above findings should be considered with great caution. 


Acknowledgement 


The authors are very grateful to Mrs. Babe Foo, Messr. Mahmud b. Sider 
and Michael Arokian for their technical help. We are also very much obliged to 
Miss Amy Leong for typing the manuscript, and to Dr. Liew Fah Seong for going 
through the manuscript and for his constructive criticisms. 


References 


Atkinson, L. R., 1973. The gametophyte and family relationships. In A. C. Jermy 
et al (Ed.) — The phylogeny and classification of the ferns, 73-90. 


Ching, R. C, 1940. On natural classification of the family Polypodiaceae. 
Sunyatsenia 5, 201-268. 


ee C., 1938. Filicinae. In F. Verdoorn (Ed.) — Manual of Pteridology. 
22-550. 


copeeesls E. B., 1947. Genera Filicum. Chronica Botanica. Waltham Mass.. 


Erdtman, G., 1957. Pollen and Spore Morphology/Plant Taxonomy — Ul. — 
Gymnospermae, Pteridophyta, Bryophyta. 


94 Gardens’ Bulletin, Singapore — XXX (1977) 


—____—_— & P. Sorsa, 1971. Pollen and Spore Morphology/Plant Taxonomy 
— IV. — Pteridophyta. 


Harris, W. F., 1955. A manual of the spores of New Zealand Pteridophyta. N.Z. 
Dept. Sci. Industr. Res. Bull, 116, 186 pp. Willington. 


Holttum, R. E. 1947. A revised classification of Leptosporangiate ferns. J. Linn. 
Soc. (Bot.) Lond. 52, 123-158. 


1949. The classification of ferns. Biol, Rey, 24, 267-295. 


1954. A Revised Flora of Malaya — II. Ferns of Malaya, 299-574. 
Govt. Printing Office Singapore. 


1973. Introduction to Pteridophyta. In J. C. Willis (Revised by 
H. K. Airy Shaw) Dict. FI. Pl. & Ferns. 8th. edition. Cambridge University 
Press. 


Lovis, J. D., 1973. A biosystematic approach to phylogenetic problems and its 
application to the Aspleniaceae. In A. C. Jermy et al (Ed.) — The phylogeny 
and classification of the ferns, 211-227. 


Manton, [., 1961. Evolution in the Pteridophyta. In P. J. Wanstall (Ed.) — A 
Darwin Centenary Bot. Soc. Brit. Isle Conf. Rep. 6, 105-119 


Mickel, J. T., 1973. The classification and phylogenetic position of the Dennstaed- 
tiaceae. In A. C. Jermy et al (Ed.) — The phylogeny and classification of 
the ferns, 135-144. 


Nayar, B. K., 1970. A phylogenetic classification of the homosporous ferns. Taxon 
19, 229-236. 


Nayar, B. K. & S. Devi, 1963. Spore morphology of some Japanese Aspidiaceae. 
Pollen Spores 5, 354-372. 


1964 a. Spore morphology of Indian ferns-I. Aspidiaceae. Grana 
Palynol. 5, 80-120. 


1964 b. Spore morphology of Indian ferns-II Aspleniaceae & 
Blechnaceae. Grana Palynol. 5, 222-242. 


1966. Spore morphology of the Pteridaceae-I. The Pteridoid ferns 
Grana Palynol. 6, 476-497. 


1967. Spore morphology of the Pteridaceae — II. The Gymno- 
grammoid ferns. Grana Palynol. 7, 568-600. 


1968a. Spore morphology of the Pteridaceae. III. The Dicksonioid, 
Dennstaedtioid and Lindsayoid ferns. Grana Palynol. 8, 185-203. 


1968b. Spore morphology of the Pteridaceae. IV. Taxonomic and 
phyletic considerations. Grana Palynol. 8, 517-535. 


Nayar B. K. & S, Kaur, 1963. Spore morphology of some Indian members of the 
Lomariopsidaceae. Pollen Spores 5, 87-94. 


Pichi Sermolli, R. E. G., 1958. The higher taxa of the Pteridophyta and their 
classification. Uppsala Univ. Arsskrift., 6, 70-90. 


1973. Historical review of the higher classification of the Filicopsida. 
In A, C. Jermy et al (Ed.) — The phylogeny and classification of the ferns, 
11-40 


Reimers, H., 1954. Pteridophytg. Farnpflanzen. In Engler’s Syllabus der Pflanzen- 
familien, 12 Aufl. I, 269-311. Berlin. 


Malayan Dennstaedtiaceae 95 


Sen, U. & T. Sen, 1973. Anatomical relationships between the Oleandra and 
Nephrolepis groups. In A. C. Jermy et al (Ed.) — The phylogeny and classifi- 
cation of the ferns, 155-172. 


Sledge, W. A., 1973. Generic and family boundaries in the Aspidiaceae and 
Athyriaceae. In A. C. Jermy et al (Ed.) — The phylogeny and classification 
of the ferns, 203-210. 


Swain, T. & G. Cooper-Driver, 1973. Biochemical systematics in the Filicopsida. 
In A. C. Jermy et al (Ed.) — The phylogeny and classification of the ferns, 
111-134. 

Tardieu-Blot, M. L., 1963a. Sur les spores de Lindsaeaceae et de Dennstaedtiaceae 
de Madagascar et des Mascareignes, étude de palynologie appliquée a la 
systematique. Pollen Spores 5, 69-86. 

1963b. Sur les spores de Pterideae Malgaches. Pollen Spores §, 
337-353. 


1965. Sur les spores d’Adiantaceae, Aspleniaceae, Thelypteridaceae 
et Athyriaceae de Madagascar. Pollen Spores 7, 319-338. 


Van Cotthem, W. R. J., 1973. Stomatal types and systematics. In A. C. Jermy 
et al (Ed.) — The phylogeny and classification of the ferns, 59-71. 


Walker, T. G., 1973. Evidence from cytology in the classification of ferns. In A. C. 
Jermy et al (Ed.) — The phylogeny and classification of the ferns, 91-110. 


Willis, J. C., 1973. Dict. Fl. Pl. & Ferns. (Revised by H. K. Airy Shaw). 8th ed. 
Cambridge Univ. Press. 


rengys soctr ew me St 


tire. Ww : p 5, 
br) “egsodibiqdAedt. b 


mincitaae; hes ais =, fbS) We > | 
nt) Muaval. $8. FG 18a, : pac. 

shieqorilT ont f Za ATONE Es an cl thee 

aS) SI 3S Onn AWB oan Ag 


“~~ 

sesoeiibssiennsO ot 35 56 saan | of enoge eal we 
; Sythe seOlOt ie Hh OUTS eos visrsuen, 
; AF CRS we waned ey 

a ( by vie ay} i): ‘eh weg 

‘gy a era 

oy bh 25% ee dir ine 


jeX aaa 


. 
- 
- 


(2 : i! us HZ. EXD} 
sae On iw Eat AR 


Plate 1. Figs. 1-3: Pteris vittata; | in the centre 


ne . . 
aii OF perine tay 


obl.e.v.; 3 — detailed structure of distal | : 
(abbrev. in legends: pr.p.v. = proximal polar view, obl.e.v. 
view, d = distal) 


Plate 2. Fig. | — Acrostichum aureum, pr.p.v.; figs. 2 & 3 — Lindsaea scandens, pr.p.v. & 
e.v. respectively; figs.-4 & 5: Isoloma divergens, 4 — pr.p.v., 5 — optical section of 
4: fis. 6 & 7: Microlepia speluncae, 6 — pr.p.v., 7 — optical section of 6; fig,.8:-= 
Orthiopteris kingii, optical section of pr.p.v. 


2 — pr.p.v.; 


Plate 3. Figs. 1 & 2: Asplenium nidus var. musifolia, | — e.v., 
Blechnum orientale, 3 — obl.pr.p.v., 4 — e.v. with broken perine; 


Dryopteris sparsa, 5 — obl.pr.p.v., 6 — e.v. 


Plate 4. Figs. 1-2: Elaphoglossum callifolium, 1 — e.v., 2 — d.p.v.; figs. 3 & 4: Heterogonium 
pinnatum, 3 — left, e.v., right, d.p.v., 4 — pr.p.v.; figs. 5 & 6: Oleandra pistillaris, 
d.p.v., 5 — focus on perine, 6 -— optical section. 


uf 


Plate 5. Figs. 1-3: Teratophyllum aculeatum var. montanum, 1 — d.p.v. showing detailed 
structure of perine, 2 — pr.p.v., 3 — obl.e.v.; figs. 4-6: Arcypteris irregularis, 
4 — obl.pr.p.v., 5 — pr.p.v. in optical section, 6 — d.p.v. 


Plate 6. Figs. 1-3: Davallia divaricata, 1 — pr.p.v. in optical section, 2 — detailed structure 
of exine in pr.p.v., 3 — e.v. in optical section; figs. 4-8: Humata heterophylla, 
4 — obl.e.v., 5 — pr.p.v., 6 — d.p.v., 7 — e.v., 8 — e.v. in optical section. 


Plate 7. Figs. | & 2: Histiopteris stipulacea, | — e.v., 2 — obl.pr.p.v.: figs. 3 & 4 Nephrolepis 
biserrata, 3 — pr.p.v., 4 — e.v.; 5 & 6: Stenochlaena palustris, 5 — pr-.p.v. in optical 
section, 6 — e.v. in optical section. 


2 — d.p.v., 
5 — pr. p.v. 


3 
’ 


ine 
ion 


1 sect 


tica 


in op 


th ruptured per 


wi 
V 


.e.V 
is, 4—e 


1 — obl 
ivalvis, 
V 


is b 


isecta 
. 7—e 


1Sec 


Hypolep 
LV 


p 


is fenu 


5 


6 — pr 


4 


figs. 


t 


ion 


ystopter 


¢ 


V 
1 sec 


1-3: 
pr.p. 
ica 


in opt 


igs 


Fi 
lees 


Plate 8 


Plate 9. Figs. 1-2: Lomagramma sumatrana, 1 — e.v., 2 — obl.pr.p.v.; fig. 3 — Sphenomeris 
chusana, obl.e.v.; fig. 4 — Tapeinidium pinnatum, obl.e.v.; figs. 5 & 6: Athyrium 


pinnatum, 5 — obl.pr.p.v., 6 — e.v. 


lata, d.p.v., 
d.p.v. 


icu 
6 


fig. 3 — Egenolfia append 
ion 


1 sect 


tica 


oa .p.v. in op 


obl.e.v., 5 — pr 


, 4 


inea insignis 


Bra 


Plate 10. Figs. 1 & 2 — Bolbitis heteroclita, obl.c.v 
figs. 4-6 


An Enumeration of the Malesian Species of Aralia L. 
by 
W.R. PHILIPSON 


Department of Botany 
University of Canterbury, New Zealand 


The species of Aralia in South-east Asia and the adjacent islands have been 
subject to considerable misunderstanding. The present account takes a broad view 
of specific limits, following along lines originally proposed by van Steenis (Bull. 
Bot. Gard. Buitenz. ser. 3, 17 (1948) 391). 


Aralia 


Linn. Sp. Pl. (1753) 273. — Acanthophora Merr. Philip. J. Sc. 13 (1918) 316; 
Steen. Bull. Bot. Gard. Btzg. ser 3, 17 (1948) 390. 


Sparingly branched shrubs or small trees, or climbing, rarely (extra Malesia) 
herbaceous, glabrous or hairy, often prickly. Leaves pinnate to tri-pinnate, usually 
with leaflets at the insertion of the lateral pinnae; leaflets serrate; petiole with a 
sheathing base. Inflorescence a terminal panicle; flowers sessile or pedicellate, with 
an articulation below the flower; calyx with 5—6 teeth; petals 5-6, imbricate; ovary 
2-6 celled; styles 2-6 free or shortly connate below. Fruit a fleshy drupe; pyrenes 
cartilaginous compressed; endosperm uniform. 


Distr. More than 30 species in North America and Asia. In Malesia 6 species, 
extending from Sumatra and the Malay Peninsula through Borneo and Celebes to 
the Philippines and West New Guinea. 


Eco. Usually on scrubby hillsides and in second growth, often in ravines or 
near streams, or in thickets near or above the limit of tree-growth. Occurring at 
low altitudes (100 m) but usually in the montane zone, up to 3000 m. 


Notes. Hui-Lin Li in Sargentia 2 (1942) 101, treats some species that extend 
into Malesia. Merrill considered that the climbing habit and recurved spines of 
Acanthophora justified its separation as a distinct genus, but more recent authors 
have not agreed. 


KEY TO THE SPECIES 


1. a. Flowers sessile (capitate) or very shortly pedicellate .....................:0.00 2 
i rowers peaicellate iCumbellate) 45.00. Lh ocean ee eeees 3 
2. a. Flowers sessile, underside of leaf + densely tomentose, hairs of the 


branches and inflorescence + appressed and felted, bracts around the 
PREM I TEN PAT oe. soaks. Tadeo eee ceca eens 1. A. dasyphylla 


b. Flowers short pedicellate, underside of leaf sparsely tomentose, hairs of the 
branches and inflorescence + patent, bracts around the capitula less densely 


I Re ed i ov cciaa « siildnweu ag gh~s omansed dp adaeee 2. A. javanica 
3. a. Climbing or scrambling liane, spines curved .................. 3. A. scandens 
b. Erect shrubs or small trees, spines straight ................cccscseseceeeeeeeeeeees 4 


98 Gardens’ Bulletin, Singapore — XXX (1977) 


4. a. Leaflets glaucous beneath, margins with few crenations, fruit small (+ 


3 MM TOME) .....00.00s00svaveenenpussvaes sys peeudbnaie sen 4. A. bipinnata 

b. Leaflets green (or with fawn pubescence) beneath, margins serrate, fruit 
rather larger (4-6 mm long):).,6..5000)s.80100. nema ey 5 

5. a. Young parts and under-surface of leaves glabrous (but with small spines) 
stebeccesbocsnsdtwbanna sec see ekg hey te MehtOA te onkhs the niuics + aitinnntna an 5. A. ferox 

b. Young parts and under-surface of leaves pubescent ............ 6. A. montana 


1. Aralia dasyphylla Mig. Fl. Ind, Bat. 1, 1 (1856) 751. — A. urticifolia BI. 
ex Mig. Ann, Mus. Bot, Lugd. -Bat. 1 (1863). — A. dasyphylla var. strigosa Miq. 
Ann. Mus. Bot. -Lugd. — Bat. 1 (1863) 9. — A. dasyphylla var, latifolia Miq. 
Ann, Mus. Bot. Lugd. -Bat. 1 (1863) 9. — A. beccarii Ridl. J. Mal. Br. R. As. 
Soc. 87 (1923) 64. — A. dasyphylla var. urticifolia (Bl. ex Miq.) Bakh. in Blumea 
6 (1947) 367. — A. dasyphylla var. typica Bakh, -Oostr, in Back. Bekn, Fl. Java, 
(em. ed.) 7 (1948) fam. 159, 18. — A. dasyphylla var. dasyphylla Back. & Bakh. 
Fl. Java, 2 (1965) 710. 


Prickly shrub or small tree, often unbranched, to about 5 m high, young parts 
densely brown pubescent. Leaves forming large rosettes at the summit of the stems, 
about 1 m long (or more), bi- or tripinnate, the petiole, rhachis and lateral 
rhachides prickly or unarmed, densely pubescent; leaflets subsessile or petiolule 
c. 5 mm long (or longer), usually densely pubescent on the lower surface, less 
dense above, ovate to oblong-ovate, c. 5-14 (18) x 3-5 (10) cm, base rounded to 
sub-cordate, apex accuminate, margin finely or sometimes coarsely serrulate. 
Inflorescence a large terminal panicle, densely brown pubescent, ultimate branches 
arranged racemosely, ending in heads of several sessile flowers. 


Distr. Malay Peninsula, Sumatra, West and Central Java. Also northwards to 
southern China. 


Eco. Primary forest and second growth in deep ravines or open hillsides, 
occurring from low altitudes (+ 100 m) to 2,500 m. 


Notes. The capitulate flowers are characteristic (see also under A. javanica). 
The presence of this species in the Malay Peninsula has often been overlooked, 
though it extends into southern China. Variability is discussed by van Steenis 
(Bull. Bot. Gard. Btzg. ser 3, 17 (1948) 391). 


2. Aralia javanica Miq. Pl. Jungh. 3 (1855) 420. 


A shrub or small tree, often unbranched, young parts covered with brown 
pubescence which persists on the stems and rhachides of the inflorescence as + 
patent hairs. Leaves tufted at the ends of the branches, bipinnate; leaflets variable 
in size, subsessile or the petiolules up to c. 2 cm long, blade ovate to elliptic, up 
to 18 X 8 cm (usually smaller), both surfaces with sparse short appressed bristly 
hairs, base cuneate to truncate, apex accuminate, margin finely and unevenly 
serrulate. Inflorescence a large terminal panicle; the ultimate branches bearing 
heads (or sub-umbellules) of c. 10 flowers, surrounded by an involucre of small 
linear bracts. 


Distr. Central and West Java (Dieng, Surakarta, G. Papandayan, Mt Malabar, 
G. Prahu). | 


Eco.. Mountain forests, 2000-3000 m. 


Notes. This imperfectly known species may prove to be a form of the wide- 
spread A. dasyphylla, from which it appears to differ in the shortly pedicellate 
flowers, the sparser leaf-tomentum, the more patent hairs on the inflorescence 
branches and the bracts of the umbellules less thickly enveloped in hairs. 


Malesian Aralia 99 


3. Aralia scandens (Merr.) Ha in Nov. Sist. Vyssh. Rast. 11 (1974) 229. — 
Acanthophora scandens Merr, in Philip. J. Sc. 13 (1918) 316. — ‘‘Aralia ferox 
Mig.” King in J. As. Soc. Beng 67 (1898) 45. 


Prickly scandent shrub, glabrous, reaching a height of 10 m or more, stems 
c. 2.5 cm thick. Leaves dispersed (+ 30 cm apart), up to 1.5 m long, tri- or 
quadri-pinnate, prickly on the petiole, rhachides, and sometimes on the leaf veins; 
leaflets ovate to elliptic-ovate or ovate-lanceolate, base rounded or sub-cordate, 
apex acuminate, margins finely spinulose-denticulate. Inflorescence a large terminal 
spiny panicle, the main rhachis to c. 60 cm long, bearing secondary branches singly 
or in whorls, up to 50 cm long; the ultimate branches 1-4 cm long, subtended by 
lanceolate bracts, racemosely arranged, ending in umbellules; umbellules 10-20 
flowered, pedicels slender 10-12 mm long. 


Distr. Malay Peninsula: Perak, Selangor, Pahang. Sabah: Mt. Kinabalu. 
Celebes: Menado, South Celebes. Philippines: Luzon, Panay, Catanduanes, 
Mindanao. 


Ecou, Thickets on mountainsides, often near streams, or among second growth. 
Between 180-1550 m altitude. 


Notes. The only species with the habit of a liane, with spaced leaves, and 
recurved spines. The flowers are whitish or yellowish, and slightly fragrant. Visited 
by numerous small bees. Fruit fleshy, purple. Scrapings of the bark are applied to 
wounds and a decoction of the boiled bark is drunk to releave internal pain 
(Mindanao). 


4. Aralia bipinnata Blanco, FI. Filip. (1837) 222. — A. hypoleuca Presl. Epim. 
(1851) 250. — “A. javanica Miq” F. Vill. Nov. App. (1880) 101. — A. glauca 
Merr. in Philip. J. Sc. 2 (1907) 291. A. apoensis Elmer, Leafl. Philip. Bot. 7 (1914) 
2325. A. bipinnata f. inermis Steen, in Bull. Bot. Gard. Btzg. ser 3, 17 (1948) 392. 


A shrub or small sparsely branched tree to 7 m high, with prickly stems, Leaf 
to 1.5 m or more long, forming large crowns at the ends of the branches, bipinnate, 
with some prickles, especially on the petiole or unarmed; leaflets sessile or with a 
short petiolule, ovate to lanceolate, apex acute or acuminate, base rounded to 
cordate, margin conspicuously crenate, upper surface green, glabrous, lower surface 
glaucous, pubescent along the veins and sometimes sparingly on the mesophyll, or 
almost glabrous throughout. Inflorescence a large terminal panicle, the peduncle 
and also usually the main branches prickly, the whole either almost glabrous or 
pubescent; tertiary branches usually 5-10 cm long, ending in umbellules, and 
bearing a small number of lateral umbellules, or branches; umbellules with many 
radiating pedicels (about 20-30); pedicels 5-10 mm long. 


Distr. Philippines: Luzon, Leyte, Negros, Mindoro, Mindanao. West New 
Guinea: Vogelkop. Also in Taiwan and Okinawa. 


Eoo.. In rather open forests, ravines, and in thickets and second-growth; 
occurring at about 700 m, but chiefly between 1000-2450 m. 


Notes. van Steenis discusses the variability in pubescence and the development 
of spines (Bull. Bot. Gard. Btzg. ser 3, 17 (1948) 392. 


5. Aralia ferox Miq. FI. Ind. Bat. 1, 1 (1856) 750. — A. filicifolia Ridl. J. Fed. 
Mal. St. Mus. 8 (1917) 42. 


Spiny shrub or small tree, usually unbranched, to about 10 m high. Leaves 
forming a large rosette at the summit of the stem, up to about 1 m long, bi- or 
tri-pinnate, prickly on the petiole, main rhachis, and often on the lateral rhachides; 
leaflets sessile or petiolules to c. 5 mm, ovate or ovate-oblong, variable in size, 
base truncate to rounded or cuneate, apex acute acuminate, margin sharply serrate, 


100 Gardens’ Bulletin, Singapore — XXX (1977) 


both surfaces with small bristle-like spines, especially on the veins, Inflorescence a 
large terminal panicle, 25-50 cm long, glabrous, tertiary branches disposed singly 
or in sub-verticils, ending in umbellules; flowers about 10-12 per umbellule. 


Distr. Sumatra: West Coast, Mt Kerintje. Java: Gedeh, Patuha and Tang- 
kuban Prahu. 


Ecot. In montane scrub and among scattered trees; occurring between 
1900-2900 m. 


forma nana Steen. in Bull, Bot, Gard. Btzg. ser 3, 17 (1948) 394, fig. 1. 


Smaller, probably 0.5-1.0 m tall, leaves tri-pinnate, 30 cm long, spiny all over, 
leaflets 4-13 X 2-7 mm, rhachides of the ultimate pinnae winged. 


Distr. Sumatra: known only from Mt Talang. 
EcoL. Growing about 2500 m altitude. 


Notes. van Steenis considers this interesting dwarf form to represent the 
extreme of a series in size variability. 


6. Aralia montana Bl. Bijdr. (1826) 870. — Panax armatus Wall. Cat (1832), 
4933 nomen nudum; G, Don Gen. Syst. 3 (1834) 386. — Aralia montana var. 
acutata Miq. Ann. Mus. Bot. Lugd. -Bat., 1 (1863) 9. — A. armata (Wall.) Seem. 
Journ. Bot. 6 (1868) 134. — A. thomsonii Seem. Journ. Bot. 6 (1868) 134. — A. 
montana var. crassifolia Bakh. — Oostr. in Back. Fl. Java (em. ed.) 7 (1948) fam. 
159, 18. 


A shrub or small tree, frequently unbranched, with prickly stems. Leaves to 
1 m or more long, forming large crowns at the ends of the branches, bipinnate, 
usually with some prickles, especially on the petiole, or unarmed. Leaflets sessile 
or with a short petiolule, ovate, up to 14 X 7 cm, apex acute to acuminate, base 
truncate or rounded, oblique in lateral leaflets, margin sharply serrate, upper surface 
with the remains of a strigose tomentum, often + rugose, lower surface often 
with a + velvety tomentum, or with more harsh hairs + confined to the veins. 
Inflorescence a large terminal panicle, the peduncle and branches tomentose, 
prickles, if any, confined to the peduncle and main rhachis; secondary branches at 
intervals along the main rhachis, c. 35 cm long, bearing numerous tertiary branches 
along their length; tertiary branches usually c. 6 cm long, ending in umbellules, 
and often bearing a number of lateral umbellules; umbellules with many radiating 
pedicels; pedicels usually 12-15 mm long, occasionally shorter, pubescent. Lobes 5, 
5-celled, glabrous. 


Distr. Malay Peninsula, Sumatra, Java, Sarawak, Sabah, Celebes. 


EcoL. Primary and secondary forest, bamboo forest and low-lying moist 
ground, from near sea level to 2600 m. 


Notes. This species is considered to include all west Malesian examples with 
pedicellate flowers and pubescent leaves, This broad concept is contrary to former 
treatments which have recognized several species (A. thomsonii, A. armata), The 
alliance with A. chinense Linn, is also very close. The application of names to this 
and other Javanese species has been very confused. The position is ably discussed 
by Koorders and Valeton Bijdr. 7 (1900) 52 et seq. and by van Steenis Bull. Bot. 
Gard. Btzg Ser 3, 17 (1948) 391. In most specimens the lower leaf surface and the 
pedicels are densely tomentose, but examples occur in which the leaf is only 
sparsely hairy and the pedicels may be glabrous. A. armata appears to be within 
the range of viaration of the complex as also does A. decaisneana Hance of China. 
The most aberrant specimens are those with glabrous umbellules which are mostly 


from Sumatra but also from Java: they may indicate that A. foliolosa Seem. should 
also be included in this complex. 


Scanning Electron Microscopical Studies on the 
Spores of Pteridophytes 


XI. The Family Oleandraceae (Oleandra, Nephrolepis and Arthropteris) 
by 
LIEW FAH SEONG 


Department of Botany 
University of Malaya 


Abstract 


The monolete bilateral spores of three genera of ferns belonging to the Oleandraceae 
(Oleandra, Nephrolepis and Arthropteris) were examined by scanning electron microscope. 
It is found that the spores of Oleandra are characterized by a large folded perine, and densely 
echinate exine bearing pointed spines or blunt supporting rods. Arthropteris possesses spores 
with more or less similar perine characteristics, and considering its morphological similarities 
with those of Oleandra, this genus may well be included in the family Oleandraceae. Spores 
of Nephrolepis, on the other hand, being characterized by insulate or verrucose perines, 
thus resemble more closely those of the Davalliaceae, the closest relative of Nephrolepis. 
However, many of its morphological and anatomical characteristics are so distinctive and 
unique that it clearly constitutes a natural group by itself, and should, as proposed by Ponce 
de Leon (1953), be treated as the only member of an independent family, the Nephrolepidaceae. 


Introduction 


The taxonomic and phylogenetic position of the fern genera Oleandra, Nephro- 
lepis, Arthropteris and Psammiosorus have long been debated and remain contro- 
versial. Oleandra Cav. is a genus of the tropics and has about 40 species. It differs 
from other fern genera in many features, and for that reason its taxonomic position 
is not fully established. Attributing great importance to the articulation of the stipe 
to the stem J. Smith (1866) first segregated Oleandra from the Aspidiaceae and 
considered it the sole genus of the new tribe of Oleandreae. After the detailed 
anatomical study of the Japanese species O. wallichii by Ogura (1938), Oleandra 
was considered as an isolated genus, and treated as the only genus in the family 
Oleandraceae (Ching, 1940). Later, many authors accepted it as a monogeneric 
family (Dickason, 1946; Ponce de Leon, 1953; Pichi-Sermolli, 1965). Others 
however, have either treated it as “a genus incertae sedis’ (Bower, 1928), or placed 
it among many other groups of ferns. Of the latter Diels (1889) arranged it in the 
independent monotypic Tribus Oleandreae of Polypodiaceae, and Christensen 
(1938) put it in the monotypic subfamily Oleandroideae of the same family. Still 
other pteridologists included it in the Dennstaedtiaceae (Holttum, 1947), in the 
Davalliaceae (Copeland, 1947; Alston, 1956; Tardiue-Blot, 1958), or in the sub- 
family Oleandroideae of Davalliaceae (Tindale, 1961; Crabbe, Jermy & Mickel, 
1975). 


The genus Nephrolepis Schott has about 30 species distributed in the 
tropics, Japan and New Zealand. Pres] (1836) included Nephrolepis in the section 
Nephrodiariae of his tribe Aspidiaeae. Most workers placed it either in the family 
Davalliaceae (Ching, 1940; Dickason, 1946; Copeland, 1947), or included it in a 
separate group placed near the Davalliaceae (Nayar, 1970; Holttum, 1971, 1973). 
However, Ponce de Leon (1953) established the family Nephrolepidaceae for this 
group of ferns. Nephrolepis is usually classified with Oleandra because “they are 


101 


102 Gardens’ Bulletin, Singapore — XXX (1977) 


probably more nearly related together than either is to any other genus” (Holttum, 
1954). 

Arthropteris J. Smith is a small genus of about 20 species and distributed in 
the Old World tropics and southwards to Australia and New Zealand. ““The species 
of Arthropteris are in utter confusion” (Copeland, 1958). The affinity of Arthrop- 
teris is not clear either (Holttum, 1966). It has certain characteristics which are 
similar in some extent, to the genera Oleandra, Nephrolepis, Davallodes, Elaphog- 
lossum, Thelypteris, Teratophyllum, Ctenitis and Tectaria, It is usually placed 
near Nephrolepis, but may not be its nearest relative (Holttum, 1966). 


The monotypic genus Psammiosorus C. Chr. of Madagascar is peculiar and 
shows features in common with Arthropteris (Copeland, 1947) and the Davallioid 
ferns (Christensen, 1932; Holttum, 1966). It is always grouped with the Davallioids 
or Oleandroids (Pichi-Sermolli, 1965), but its actual taxonomic position and 
affinities are not clear. 


These four genera have in the past and recently been variously grouped 
(Holttum, 1966, 1971; Nayar, 1970; Sen & Sen, 1973; Crabbe, Jermy & Mickel, 
1975): in one family, the’ Oleandraceae; in two separate families, the 
Oleandraceae (Oleandra) and Davalliaceae (Nephrolepis, Arthropteris and 
Psammiosorus) or in more than two families (Oleandraceae, Nephrolepidaceae 
and Davalliaceae). In view of the controversy regarding the taxonomic problems 
and phylogenetic relationships of these genera, it was thought that a detailed study 
of them is a project worthwhile to undertake. 


Spores of ferns are becoming increasingly important in fern taxonomy (Nayar, 
1964; Wood, 1973; Moe 1974; Liew, 1976c). Because they offer both reliable and 
stable morphological characters in (i) distinguishing species in some genera, (ii) 
in differentiating genera, subgenera and characterizing families (Brown, 1960; Liew, 
1976b) and (iii) in the tracing of possible phylogenetic relationships and trends of 
evolution between taxa (Brown, 1960; Sorsa, 1964; Nayar & Dewi, 1968; Wagner, 
1974) the present author decided to investigate this character and use it as a means 
to tackle the problems raised above. Scanning electron microscopical observation 
is preferred and undertaken because it offers the combined superiority of greater 
depth of field, high resolution, and visually presents three dimensional topographical 
information which reveals subtle and spectacular structures of spore surfaces 
previously unobtainable with the optical microscope. 


Materials and Methods 


Spores of the three genera of ferns belonging to the Oleandraceae (Oleandra, 
Nephrolepis and Arthropteris) were taken either from fresh plant material collected 
in the fields or from herbarium sheets deposited in many institutions (NY, SING, 
TAI, KLU and HK; see Table I for details). Fresh fronds were carefully dusted 
for their spores and spores from herbarium specimens were obtained with the use 
of clean tooth picks. 


Before making scanning electron microscopical observation spores of each 
species are dispersed in a drop or two of Hoyer’s solution (Anderson, 1954) and 
checked preliminary for their proper identity under an ordinary light microscope. 
Details of the method of preparation and observation on scanning electron micro- 
scope were as described before (Liew, 1976a). Many models of scanning scopes 
were employed in the present investigation, e.g., JOEL JSM 15, JSM U3, Cambridge 
S4-10, and Hitachi MSM 4. Black and white photomicrographs were recorded on 
ordinary 120 negative or on Polaroid films with (105 P/N, 55 P/N) or without 
(42, 107) negatives. Sizes of spores were measured either in polar x equatorial axes 
(P x E) or in polar x longest equatorial x shortest equatorial axes (P x El x E2). 
Values given are averages of five to ten readings. 


‘gqetiva Joye sarods Jo sinjdjnd¢ 9 ‘sedpls jNnoYyIM soe VoVJING ¢ ‘saloeds pupuvajQ Jo serods oy} 0} IvjIUIs Ivodde sorodg y 


on ‘sospll posivl SUIWIOJ oWOs pue ‘eUNdE UT Aj[eIoodso ‘oures “|Z ‘BuIpnyjoid pue yuoulWoId AIDA 
S __SI[NAOQN} IO BVONIIBA “OZ div] “oJEINSUT BVONIIOA JO SOBPII “G] ‘[[PUIS ‘d}e[NSUT S¥ONIIOA JO sodpll ‘g] ‘OLONIIOA JO SOBPLI poOoUUOSID\UI ‘peosg 
‘oye[NSUT YM “Z] ‘usdo syovso Ajises “g] ‘YO poysnoys ApIseo “c] ‘({[e1 ™ pO ¢ 0} dn) sqnppo yunIq “py ‘poytod ‘(7 ¢ oj dn) 3uo; soulds “¢|[ ‘payutod 
‘Woys soutds 7] ‘sopoy Auew YM “[] “(sduIpjoy Ivjndoi Jo/%» sodpis posted) soinjonsys Ie[NZo11 JO snoydiowe YUM ‘oO ‘spjoy Aueul YUM “6 “WJoUs 
‘Tews sassoooid “g ‘popunos ‘jJoys sassedoid ‘1 ‘sassodo0id YIM “9 ‘(suodhjod oB1e] YIM) a3eYdO]T *¢ ‘YJOOUIS “p ‘(SoeJINS) YSnoI “¢ “UIy) “7 *yoIy} “| 
‘IOAR] OUIXD ‘p ‘JoART IeUWM]OS JoUUT ‘oUeS ‘9 ‘IOAR] JoyNO ‘ouIes “g ‘oULIOd ‘D ¢ 
‘sIxe [ellojyenbg xX sIxe Iv[og = AXdz 
‘soutddijiyg “Yq ‘vourny 
MON ‘ON ‘ieosedepey] “al ‘vAvyeyl “VA, ‘uvureH ‘ORY ‘Tremeyy ‘HY SesouNIO,y ‘ey ‘ft.7 ‘Iq ‘eidoryyg “q@ feory eIsOoD “WO ‘OoulOg ‘Og ‘vijelsny ‘wy 
9bP “hI ‘€1 10/% 719 be-LE X €€-S7 (IVD) LI€9 OFM “LW :ea seid (HH) WYyoNeEMm “67 
bP ‘P19 ‘1% ‘ZI ‘Sq Le-0e X [E°9¢ (AN) 19007 9INIDOW “V ‘A :0H suryD (PITA) BeENpun “47 
bP ‘pI? ‘ZI ‘119 SS-Sb X 8E-1E€ (ATM) 181 UlYO ‘O'S cB ‘IYO 'O (Msg) sueyysid § “¢7Z 
bP ‘pI JO ZI9 ‘172 ‘ZI ‘TT ‘Sq Sb-8E X 9€-0€ (IVL) OL6I ‘91 ‘AON “u's ‘zovusH “yg “Ud "ABD SIULIOJNIOU “ZZ 
PP ‘€19 ‘II ‘8q Ov-0E X OE-SZ (NT) €88L 2491S ‘OD ‘gq “AW *‘deuog evolieosesepeul *|Z 
SoT[IUBARD D4pudIaIO 
PP ‘81 ‘74 | | Te-Sc X C@C-LI (AT OSs yeWeg “y :eW uyny (UlIng) suesIpel “OZ 
¢ yP_‘€ ‘74 8E-97 X ST7-0C (AN) $89 J9sorJ “TW  J9dINg ‘DO “MA :UD HoYyIS (PITIAA) BeUNded = 6] 
61 “€ Apysis ‘7q ZE-97 X I7-LI (ONIS) 6161 “u's ‘uayNg :- ON sly WySeqioyne] “g] 
bP ‘81 ‘74 S€-8Z X S781 (IV) 0906 M2] ‘S ‘a Bq [scerg (S104) eNINsNY “Z| 
bP ‘LI ‘€ ‘74 8Z-Pe X CTBT (ONIS) 0786 WNWOH “A “A :BA ‘IYO “DO CAeD) ByeOTeJ 9] 
bP ‘LI ‘€ ‘79 LE-O€ X 67-E7 (AN) W80IZ S9qiog ‘N ‘OD ‘TH noysg (J) eyeWexo “CT 
LI ‘€ (74 ££-6¢ X €7-07 (NTD S68S 94S ‘D ‘A :FIW YD Seploluosyxsip “P| 
LI ‘€ Apysis 10 ¢ ‘Zq ZEST X LZ-1Z (QTM) prZL 2001g ‘DO “g :BI ‘ozy (MS) SOPIONfeaep “¢] 
Ov owlos “LT “bp Jo ¢€ ‘7q O£€-EZ X $7-02 (QTM) 6€8 Neuny “vy “A :eW [seid (“T) elosips0d “71 
LI ‘€ ‘7 67-ET X H7-8I (IV.L) L61IOI MT “§ “y ceg noyss (‘MS) BIVIIOSIQ. *T] 
LI ‘€ ‘79 €€-Le X ¥e-0¢ (ONIS) $LSZ UPITV-YHOMse|OW “DO “A ‘A ‘BA slIyD (Asoq) eBypoynnoe “O] 
yWoysSg sidajosydan 
bP ‘91 ‘OT ‘Eq ‘12 pS-Ip X Eb-l€ (ONIS) SSI puvjedoD ‘gq “A ‘Wd uMyJOH (IPA) Wuojseyjom *6 
© bP “€1 “Il “O14 ‘Ie S9-0S X 96-Sh (AN) SP6I “URL “U's ‘sUSUIITD “§ “W iV ‘Wg “f (‘ySIOJ) eTJOUD *g 
S P ‘ST ‘21 SwOS ‘Q1q ‘Te 8S-Sb X pb-be (ONIS) 89667 SUSWITD “S "WP ‘f 208 ‘IYO 'O (‘yourg) susdor “1 
o bP ‘SI ‘TT ‘OL ‘€9 ‘Te €S-Sp X 6€-0€ (IVL) I€LE O&M “LW cea uoysTy (‘Aseq) mMosted “9 
4 bP ‘ST ‘IT ‘OI ‘8 ‘€q ‘Te p9-Ep X OS-9E (AN) €8€LZ SUSWIIID “S "WF “f 208 ‘wg “f Clq “Y) eIeIONIGO “¢ 
3 PP ‘91 ‘ST “6 ‘bq ‘Te OS-Iv X 9b-8E (QTD 1829 Wlowseg-tystg “OD “A UH ‘TWD °O (Aewopieg) edivoouou 'y 
$ bP ‘SI “9 ‘€q +12 tS-Ob X 8€-87 ((ONIS) LE6I ‘§ “Ing “u's ‘spuounwig "A "H -DON _ WaA'y'a epodoysijop € 
= bP ‘SI ‘IT ‘€9 <18 SS-Iv X Ob-cE (ONIS) 076€d P19"ISU0D “YA ?W “NOW (YOOH) HWeTyx90q Ks 
° y bP ‘IT ‘L ‘Sq ‘Ie IS-Ob X 9E-87 (AN) 1989 WWE ‘D 'V * ‘IYO ‘O (yoelg) eyejnonsze “| 
8 | ywwg “¢ stuajdosyisp 
wn Soe = = £ Se). «Slee ee ey ree? ea re 2 a a. % 
. ¢ SINS149}904045 Wu unupgiax] pu saquiny =e 
& audjnog z(4Xd) 218 ‘40122109 ‘;4I1}090] = 
. ~D a 


‘sonsiojoerego o10ds Joy} JO SUIOS YIM JOYIOSO} UONLSIVSoAUT yUISoId OY) UT PoIpNys sUJOJ PloIpuesO JO exe], ‘[ ATAV] 


104 Gardens’ Bulletin, Singapore — XXX (1977) 


Results 


The sizes and general characteristics of the spores of all the species of these 
Oleandroid genera studied in the present investigation are given in Table I. 


The insulate or verrucose spores of Nephrolepis clearly form a natural group 
by themselves (Plates I and IL). These ellipsoid bilateral spores are monolete and 
mostly concavo-convex in lateral view. A few of them are plano-convex. The 
average size of the spore measures about 17-29 x 23-38 x 14-29 yw. Under the 
SEM, a broken outer sporoderm will reveal the inner exine layer of the spore, 
which is psilate, slightly scabrate or undulating (Plate I, fig. 8). The laesura is 
slightly raised, thin and with a short ridge in some species (e.g., in N. pectinata and 
N. davallioides) or thick and forming long and rough ridges in others (e.g. in 
N. acutifolia and N. biserrata). The sculptine or sporoderm has on its surface 
elements of various sizes, from small grains or humps to large tubercules, or with 
irregular excrescences or raised ridges of various sizes. 


The ellipsoid spores of Oleandra are totally different from Nephrolepis (Plates 
III and IV). They are monolete bilateral spores, and are plano-convex to slightly 
biconvex in lateral view. The amb is oblong in shape, All the spores have a thick, 
prominent, loose and wrinkled perine which anastomoses to form regular or 
irregular polygons or lophate sculptures. Depending upon the species concerned, 
the perine may have spinulose, blunt or dentate excrescences. The size of the spore 
is 25-42 x 30-55 x 26-32 yw. Scanning electron micrographs of broken sporoderms 
reveal that the outer layers are built on stilts, and are supported by numerous 
rods connected to the layer(s) down below. These micrographs also reveal the 
smoothness of the inner exine in many cases (Plate IV, fig. 40). Spores of Oleandra 
may differ in sculptural details, even within the same species, but they nevertheless 
form a group with a unique and distinctive general pattern. 


Spores of Arthropteris have a thick perine of various sculpturing ornamenta- 
tions (Plates V and VI). Foldings may be high or low in profile, and are covered 
with minute projections or interlacing networks. Some Arthropteris spores (e.g. 
A. articulata) closely resemble those of Oleandra species. The outer perine of most 
of the spores of Arthropteris sloughs off easily, leaving behind the naked exine with 
smooth surface and laesura. The size of the spores range 28-56 x 40-65 x 25-30 un. 
It is ellipsoid and sometimes roundish in polar view, and plano-convex to concavo- 
convex in lateral view. 


Discussion 


I. Nephrolepis 


From our studies on eleven taxa of Nephrolepis gathered from various parts of 
the world it is demonstrated that the bilateral spores of Nephrolepis species are 
rather uniform and form a clear-cut group by themselves. They have thin perine - 
and verrucose sculptine with elevations of various sizes, from slightly raised ridges — 
to conspicuous tubercules. 


In the literature, spores of many species of Nephrolepis have been studied. 
Holttum (1954) mentioned that spores of Nephrolepis have slightly and irregularly 
roughened surfaces, and are without distinctive markings. Hannig (1911) pointed 
out that “‘a perispore is unlikely to occur” in Nephrolepis (N. exaltata). This is 
contradicted here as shown in Plate I, fig. 8, which clearly shows the presence of an 
outer layer. Among the many authors who have examined the spores of Nephrolepis 
exaltata (L.) Schott was Selling (1946) who investigated Hawaiian species and 
described it as having very irregular warts and short, irregular ridges, with a size 
of 24-28 x 41-50 yu. Earlier workers have similarly investigated the spores of this 
species but did not mentioned the sculpture patterns. Marquesan (southeastern 


Spores of Oleandraceae 105 


Polynesia) spores of this taxon are + 34 w in length, and with the surface 
conspicuously tuberculate (Brown & Brown, 1931). Erdtman and Sorsa (1971) 
described the spores from British West Indies as convex-plane, 22 x 37 x 23 u 
(perine not included) with the laesura about 21 «, and the perine similar to that 
in N. biserrata. The exine stratification is obscure. Spores from North America 
(Florida) are subpsilate to tuberose, ellipsoid, monolete, and measure 38-55 x 
21-31 » (Kremp & Kawasaki, 1972). 


Harris (1955) described the spores of the New Zealand Nephrolepis cordifolia 
as having verrucate surfaces with elongated projections and of 18-40 xu. Spores of 
the same species in India (Darjeeling) are reported to be bilateral and possess a 
very thin perine forming small humps (Sen & Sen, 1973). 


Some other species of Nephrolepis have also been described by Erdtman and 
Sorsa (1971). For example, the spores from Java of N. davallioides (labelled as 
N. acuminata) are provided with a more or less verrucose, dark brownish, cf. 
perinous layer ca 1.0 » broad. The spores are reported to be concavo-convex and 
of 17 x 34 x 23 » in size (perine not included). The Ceylonese species of N. 
biserrata has convex-concave spores and measured 23 x 39 x 26 » (perine included). 
It has a perine ca 1.5 » thick, is verrucose; warts are dark brownish ca 1.3 x 
(2.0-6.0) » high. The Japanese species of N. falcata has a more or less perinous 
thickening which is columnar, wart-like and about (3.0-4.0) x (4.0-7.0) yu. The 
spores are 30 x 45 x 35 uw (perine not included). The Ceylonese species of N. 
hirsutula has convex-plane or slightly biconvex spores, 21 x 35 x 25 » thick and 
finely verrucose. 


The bilateral and more or less verrucate sculptine patterns of the spores is 
similar, if not identical, in all species of Nephrolepis so far examined, and thus 
form a rather homogeneous group by themselves. Though a few of the Nephrolepis 
species are remarkably variable, poorly defined or with bizarre features, the 
uniformity and uniqueness in their vegetative characters, however, has clearly 
demonstrated the naturalness of the whole group (Pichi-Sermolli, 1965). “It 
therefore constitutes a clearly natural genus’ (Copeland, 1958). 


Traditionally Nephrolepis has been associated with the Davallioids, Dick- 
sonioids or the Aspidioid ferns. Copeland (1947) commented that this genus has 
many morphological and anatomical characters in common with the Davalliaceae 
and affinity seems certain, but many are not close. In chromosome number 
(n= 41), scales (peltate) and indusium (reniform), Nephrolepis resembles 
Oleandra; but in many other characters, such as habit, anatomy of the stem and 
stipe, articulation of the stipe, the feature of the rhizome, venation, position of the 
sori and structure of the spores, they differ considerably (Pichi-Sermolli, 1965). 
However, as a group, this genus has spores similar to its closest relative of 
Davalliaceae ferns, Davallia, Humata, Araiostegia, Leucosteiga and Davallodes, 
all of which possess spores with markings of varying degree. Undoubtedly, the 
phylogenetic position of Nephrolepis is thus in the Davalliales. 


Since features of some Nephrolepis species are similar to some of the fern 
genera and families, especially members of the Davalliaceae, and have in the mean- 
time shown to be distinctive in other important characters, it might be better to 
group all the species of Nephrolepis in a separate and distinctive group, the family 
Nephrolepidaceae, as was suggested by Ponce de Leon (1935). The phylogenetic 
affinity of this family is with the family Davalliaceae. 


Il. Oleandra 


The bilateral spores of the genus Oleandra are also homogenous in having a 
distinctive, thick, smooth or wrinkled perine with a few or many protruding folds. 
The folds may form an irregular network having large meshes (lophate). Spinulose 
excrescences distributed on the surface may be few or numerous. Usually these 


106 Gardens’ Bulletin, Singapore — XXX (1977) 


are broader and thicken at the base tapering to sharp apices. Some of these 
structures have blunt rods. 


In the literature, all workers who have done palynological work on Oleandra 
have reported similar findings. For example, Erdtman and Sorsa (1971) described 
the convex-plane spores of O. musifolia of Ceylon as being 28 x 40 x 30 p 
(perine not included), the perine with folds of ca 5.0-10.0 » high, and anastomosing 
to form irregular polygons with broadly conical and blunt crests. The perine is 
sparsely spinulose or dentate. It is provided with densely spaced, intra-perinous 
rods about (2.0-5.0) x (0.5-1.0) » and spaced ca 1.0-2.0 p» apart. As described 
by Kremp and Kawasaki (1971) the ellipsoid spores of the Javanese Oleandra 
neriiformis measure 43-63 x 36-49 », and are monolete and perisporate or psilate 
(in this case, presumably the perine has sloughed off!). Erdtman (1957) also 
illustrated the spore of this species, Copeland (1947) mentioned that the spores 
are bilateral or angular by shrinkage of the epispore and are apparently smooth. 
Sen and Sen (1973) illustrated and described the spores of Indian species of 
O. wallichii as monolete, bilateral plano-convex in lateral view, and oblong in polar 
view. The perine is said to be folded and spiny. 


The habit of growth of Oleandra is almost unique and has been described as 
the only shrubby fern, although all species included in it are not necessary shrubby 
(Nayar, Bajpai & Chandra, 1968). The genus also shows homogeneity in many 
features, such as spore sculpture, etc. It differs from all other ferns in the anatomy 
of the stem, the position of the sori, the presence of peculiar aerial roots, and also 
in the structure of the spores (Pichi-Sermolli, 1965; Braggio, 1966). There are 
thus enough justification to treat Oleandra as an isolated genus in the independent 
family Oleandraceae Ching ex Pic. Ser., as agreed by many pteridologists (J. Smith, 
1866; Diels, 1889; Christensen, 1938; Ching, 1940). 

The ancestry of Oleandra is unknown (Copeland, 1947). Its remarkable 
resemblance in many features with some species of Davallia and Humata suggests 
clearly its close affinity with the Davallioid ferns. Nowadays, most workers place 
it under the Davalliales. 


In ferns, bilateral perinate spores are characteristic of the Aspidioids, 
Asplenioids, Lomariopsidoids and the Thelypterioid ferns, and the presence of 
spinulose excrescences over the perine is found especially common among the 
Aspidioid genera (Nayar & Devi, 1964; Nayar & Kaur, 1965). When the characters 
of the spores are taken into consideration with anatomical, morphological and 
cytological features, it is suggested that this unique group of ferns has a closer 
phylogenetic relationship with the Aspidioid ferns, as has been suggested by many 
investigators such as Presl (1836), Dickason (1946), Nayar, Bajpal and Chandra 
(1968), and Pichi-Sermolli (1965). The last author has also emphasized that its 
affinity in the second place is with the Polypodiales, rather than with the 
Dicksoniales to which affinity was alluded by most authors in the past. 


Ill. Arthropteris 


Copeland (1947) referred to the spores of Arthropteris as oblong and with 
epispore. In his study on the genus of Arthropteris in Malesia, Holttum (1966) 
mentioned that the spores of this genus have folded perispores. 


However, there are some differences of opinion concerning the sculptine 


structures even within the same species. For example, Harris (1955) reported the 
New Zealand species of A. tenella as having bilateral, winged spores with cristate 
perispore separated from the spore wall and both of these structures differ in 
contour. The surface is striate-scabrate and with a few crests. The crests, when 
seen in oblique or transverse view, form irregular, almost spinulose projections, 
9-14 » in height. The contour or outline is deeply resected. However, spores of 


ee 


Spores of Oleandraceae 107 


the same species from the same country were reported by Sen and Sen (1973) to 
be monolete-bilateral and possessing a thin and granulose (in the description) or 
verrucose (in the diagram) perine. Furthermore, Kremp and Kawasaki (1972) 
reported this same species to have monolete ellipsoid spores of 34-62 x 27-43 xu, 
and having a thin perine with psilate surface. Our SEM investigation on 4. 
tenella, however, confirmed the correct observation and description of Harris 
(1955). The verrucose spores of Sen and Sen (1973) were most probably due to 
contamination or observation on abnormal spores. Those observations by Kremp 
and Kawasaki (1972) could have been either too immature or mature spores with 
ablated perine, a phenomenon frequent among many Arthropteris species. 


Spores of several other species of Arthropteris have been studied e.g. A. 
altescandens of Juan Fernandez was reported to have spores with short elliptic to 
subcircular amb and as having a very thin (ca 0.3 «) and folded perine (folds up 
to 10 » high). Those of A. obliterata (labelled as Nephrolepis ramosa Moore) of 
Ceylon was said to have no perine, were biconvex and 39 x 55 x 40 «. The psilate 
spores of A. monocarpa of Africa and Madagascar and the spores with a large 
and folded perine and smooth exine of the African (Congo) A. orientalis have 
also been studied by Braggio (1966). 


The affinity of Arthropteris is not clear. Copeland (1947) placed it tentatively 
in the Oleandra-Davallia series. Collective similarities in aspects of frond structure, 
articulation of pinnae, terminal sori on veins, are evidence of affinity with the 
Nephrolepis. Holttum (1966) does not think that Nephrolepis is its nearest relative 
despite the placement near Arthropteris. He, however, agreed with J. Smith in 
accepting Arthropteris as a natural genus, and believed that its peculiar combina- 
tion of characters may give significant hints as to interrelationships among various 
groups of ferns. But, he is not yet clear as to how such a relationship may best be 
formally expressed in a scheme of classification. From studies on the spores of two 
Arthropteris species of Africa and Madagascar, Braggio (1966) emphasized that 
Arthropteris differs from both Oleandraceae and Davalliaceae, and that it ought 
to be referred to the order of Aspidiales because of its spore characteristics (with 
psilate exine and a wide and thin perine, with or without ornamention). 


From our observation on spore structure Arthropteris species is closer to the 
Oleandroids rather than to the Nephrolepis and Davalliaceae. This is especially 
evident from the spores of A. articulata which are almost indistinguishable from 
those of some Oleandra species (Plate V, figs. 46-49). Spores of other species of 
Arthropteris may not be more distant as they possess a smooth exine and a thick 
and variously folded perine with different sculptured patterns (Plate VI). However, 
they still display numerous structural features in common with the Oleandra 
spores. Pichi-Sermolli (1965) also pointed out the similarities of both genera, such 
as the articulation of the stipe, the shape of the sorus and indusium, the presence 
of a wide perine, and the chromosome number. When everything is taken into 
consideration, the disposition of Arthropteris in the Davalliales is undoubtedly 
natural. 

The proper systematic position of the peculiar monotypic genus Psamyniosorus 
(endemic in Madagascar) is not fully established (Christensen, 1932; Copeland, 
1947; Tardieu-Blot, 1964; Pichi-Sermolli, 1965; Braggio, 1966; Holttum 1966). 


In conclusion, the family Davalliaceae sensu Copeland (1947) can now be 
envisaged as the order Davalliales which comprises three natural groups, the family 
Davalliaceae s.s, containing more strictly Davallioid genera, the family Oleandra- 
ceae, with the genera Oleandra and Arthropteris, and the family Nephrolepidaceae, 
with the monotypic genus Nephrolepis. The proper phylogenetic position of 
Psammiosorus cannot be proposed until further study. At the level of the Order, 
the affinity of the Davalliales is close to the Aspidiales. 


108 Gardens’ Bulletin, Singapore — XXX (1977) 
Acknowledgements 


The author gratefully acknowledges his indebtedness to all directors and 
curators in charge of institutions for making herbarium materials available for the 
present investigation (NY, SING, TAI, KLU & HK). He also thanks Mr. and Mrs. 
C. S. Wang and C. F. Lo; Miss S. C. Wang, Mr. Y. C. Huang and Mr. K. W. Ho 
for their cooperation and technical assistance in the use of different models of 
SEMs at the various institutions and universities. 


Literature Cited 


Alston, A. H. G. 1956. The subdivision of the Polypodiaceae. Taxon, 5: 23-25 


Anderson, L. E. 1954, Hoyer’s solution as a permanent mounting medium. 
Bryologist, 57: 242-244. 


Bower, F. O. 1928. The Ferns (Filicales). Vol. If1. The Leptosporangiate Ferns. 
Cambridge Univ. Press, Cambridge. 


Braggio, G. 1966. Morfoligia delle spore sistematica delle Davalliales. Webbia, 
21: 725-767. 


Brown, C. A. 1960. What is the role of spores in fern taxonomy? Amer. Fern. J., 
50: 6-14. 

Brown, E. D. W. & F. B. H. Brown. 1931. Flora of Southeastern Polynesia. 
II. Pteridophytes. Bishop Mus. Bull, 89. 


Case, I. M. 1932. The development of the sorus in some species of Nephrolepis 
with some observations on points of anatomical interest. Trans, Roy. Soc. 
Edinb., 57: 258-276. 


Ching, R. C. 1940. On the natural classification of the family Polypodiaceae. 
Sunyatsenia, 5: 201-268. 


Christensen, C. 1932. The Pteridophyta of Madagascar. p. 70-74. Hayerup’s 
Boghandel, Kobenhavn. 


1938. Filicinae, In F. Verdoorn (ed.), Manual of Pteridology. p. 
522-550. Nijhoff, the. Hague. 


Copeland, E. B. 1947. Genera Filicum. Chronical Botanica, Waltham, Mass. 


1958. Fern Flora of the Philippines. Vol. I. Nat. Inst. Sci. Tech., 
Manila. 


Crabbe, J. A., A. C. Jermy & J. T. Mickel. 1975. A new generic sequence for the 
pteridophyte herbarium. Fern Gaz., 11: 141-162. 


Dickason, F. G. 1946, A phylogenetic study of the ferns of Burma. Ohio J. Sci., 
46: 73-108. 


Diels, L. 1899. Polypodiaceae. In A. Engler & K. Prantl, Die Naturlichen Pflanzen- 
familien, I (4): 139-339. Engelmann, Leipzig. 


Erdtman, G. 1957. Pollen and Spore Morphology/Plant Taxonomy. (Corrected 
reprint 1972) Hafner Pub. Co., New York. 


Erdtman, G. & P. Sorsa. 1971. Pollen and Spore Morphology/Plant. Taxonomy. 
IV. Pteridophyta. Almqvist & Wiksell, Stockholm. 


Hannig, E. 1911. Uber das Vorkommen von Perisporien bei den Filicinen nebst 
ge gi uber die systematische Bedeutung derselben. Flora, 103: 
321-346. 


ae eS 


Spores of Oleandraceae 109 


Harris, W. F. 1955. A manual of the spores of New Zealand Pteridophyta. New 
Zealand Dept. Sci. Ind. Res. Bull. 116: 1-186. 


Holttum, R. E. 1947. A revised classification of leptosporangiate ferns. Bot. J. 
Linn. Soc., 53: 123-158. 


1949. Classification of ferns. Biol. Rev., 24 267-296. 


1954. A Revised Flora of Malaya. Vol. Il. Ferns. (2nd ed., 1968) 
Government Printing Office, Singapore. 


—_—_———— 1966. The genus Arthropteris J. Sm. in Malesia. Blumea, 
14: 225-229. 


1971. The family names of ferns. Taxon, 20: 527-531. 


1973. Introduction to Pteridophyta (and) Family names of 
Pteridophyta (+ all pteridophyte entries). In J. C. Willis, A Dictionary of 
the Flowering Plants and Ferns, 8th ed. Revised by H. K. Airy Shaw. 
Cambridge Univ. Press, Cambridge. 


Kremp, G. O. W. & T. Kawasaki. 1972. The Spores of the Pteridophytes. Hirokawa 
Book Co., Tokyo. 


Liew, F. S. 1976a. Scanning electron microscopical studies on the spores of 
Pteridophytes. I. The genus Pyrrosia Mirbel (Polypodiaceae) found in Taiwan. 
Quart. J. Taiwan Mus., 24: 191-216. 


1976b. A scanning electron microscopical study on the spores of 
Lycopodium found in Taiwan. Sci. Agr., 24: 162-172. 


—_—_ 1976c. Scanning electron microscopical studies on the spores of 
pteridophytes. VII. The family Plagiogyriaceae, Taiwania, 21: 37-49. 


Moe, D. 1974. Identification key for trilete microspores of Fennoscandian 
Pteridophyta. Grana Palyn., 14: 132-142. 


Nayar, B. K. 1964. Palynology of modern Pteridophyta. In B. K. Nair (ed.), 
Advances in Palynology, p. 100-141. United Block Printers, Lucknow. 


1970. Classification of homosporous ferns. Taxon, 19: 229-236. 


Nayar, B. K. & S. Devi. 1964. Spore morphology of Indian ferns. I. Aspidiaceae. 
Grana Palyn., 5: 80-120. 


1968. Spore morphology of the Pteridaceae. IV. Taxonomic and 
phyletic considerations. Grana Palyn., 8: 517-535. 


Nayar, B. K. & S. Kaur, 1965. Spores morphology of the Lomariopsidaceae. 
J. Palyn., 1: 10-26. 


Nayar, B. K., N. Bajpai & S. Chandra. 1968. Contributions to the morphology of 
the fern genus Oleandra. Bot. J. Linn. Soc., 60: 265-282. 


Ogura, Y. 1938. Anatomy and morphology of Oleandra wallichii (Hk.) Presl. 
Jap. J. Bot., 9: 193-211. 


Perez Arbelaez, E. 1928. Die naturliche Gruppe der Davalliaceen (Sm.) Kaulf. Bot. 
Abhandl. K. Geobel, 14:1-96. 


Pichi-Sermolli, R. E. G. 1959, Pteridophyta. In W. B. Turril (ed.), Vistas in 
Botany, p. 421-493. Pergamon Press, London. 


—————. 1965. Adumbratio Florae Aethiopicae. II. Oleandraceae. Webbia, 
20: 745-769. 


110 Gardens’ Bulletin, Singapore — XXX (1977) 


1970. A provisional catalogue of the family names of living 
pteridophytes. Webbia, 25: 219-297. 
Pokrowskaila, I. M. 1958. Analyse Pollingue. Ann. Serv. Inform. Geol. B.R.G.G.M., 
24: 1-435. 
Ponce de Leon, A. 1953. Nociones de botanica Sistematica. VIII. Division 
Pteridophyta. Revta Soc. Cuban Bot., 10: 37-45. 
Presl, C. B. 1836. Tentamen Pteridographiae. Haase, Prague. 


Selling, O. H. 1946. Studies in Hawaiian pollen statistics, Part I. The spores of the 
Hawaiian pteridophytes. Bishop Mus. Spec. Pub, 37: 1-87. 


Sen, T., U. Sen & R. E. Holttum. 1972. Morphology and anatomy of the genus 
Davallia, Araiostegia and Davallodes, with a discussion on their affinities. 
Kew Bull., 27: 217-243. 


Sen. U. & T. Sen, 1973. Anatomical relationships between the Oleandra and 
Nephrolepis groups. Bot. J. Linn. Soc. 67, Suppl. 1: 155-172. 


Smith, J. 1866. Ferns, British and Foreign. p. 73. London. 


Sorsa, P. 1964. Studies of the spore morphology of Fennoscandian fern species. 
Ann. Bot. Fenn., 1: 179-201. 


Tardieu-Blot, M. L. 1958. Polypodiacees (sensu lato). Dennstaedtiacees. Aspidia- 
cees. In H. Humbert (ed), Flore de Madagascar et des Comores (Plantes 
Vasculaires). Fam. 5. 1: 1-391. Typographie Firmin-Didot, Paris. 


1964. Sur les spores de Davalliaceae et Vittariaceae Malgaches. 
Pollen Spores, 6: 537-544. 


Tindale, M. D. 1961. Pteridophyta of South Eastern Australia. Davalliaceae. 
Contr. N.S. Wales Nat. Herb. Fl. Ser. 208-211: 1-4. 


Wagner, W. H., Jr. 1974. Structure of spores in relation to fern phylogeny. 
Ann. Missouri Bot. Gard., 61: 332-353. 


Walker, J. W. & J. A. Doyle. 1975. The bases of Angiosperm phylogeny: Palynology. 
Ann. Missouri Bot. Gard., 62: 664-723. 


Wood, C. C. 1973. Spore variation in the Thelypteridaceae. Bot. J. Linn. Soc. 67. 


Suppl. 1: 191-202. 


Scale for figures of Plates I to VI 


For values of magnifications in the legend. / cm on the micrographs approximates to:— 


67u (x 150) IS5u (x 750) 6.7u (x 1500) 
40u (x 250) 10x (x 1000) 4u (x 2500) 
20u (x 500) 8u (x 1250) 2.5u (x 4000) 


1 


WG QE ‘ 
: Sk TAM 
. WS \ x S 


S 
S WG NS S y . \ XC 


SSS 
NSS 


NG 


SAY 


Plate I. Spore characteristics of Nephrolepis. 


Figs. 7: Group of spores shown at different views: 2, 3, 4: typical bilateral spores of 
Nephrolepis, with surfaces more or less smooth (2) and rough (3, 4); 5, 6: spores in polar 
view, showing laesurae thin (5) and thick (6); 7-/0: close-up views of the spore surfaces, 
showing more or less smooth (7), moderately roughened (8, 9) and rough (/0) surfaces. Note 
that 8 is of a broken spore of N. exaltata, showing the outer rough surface (perine) and the 
inner smooth surface (exine). 


N. pectinata: figs. J] (x 750), 2 (x 1000), 7 (x 2500); N. cordifolia: 3 (x 1500), 5 (x 1500); 


N. hirsutula: 4 (x 1250), 9 (x 2500), 7/0 (x 5000); N. biserrata: 6 (x 1500); N. exaltata: 8 
(x 2500). 


22 


Plate II. Some representative spores of Nephrolepis species. 


N. acutifolia: figs. 1] (x 1250), 12 (x 500); N. biserrata: 13 (x 1500); N. cordifolia: 14 
(x 1500), 75 (x 1250); N. davallioides: 16 (x 1250); N. dicksonioides: 17 (x 1250); N. exaltata: 
18 (x 1000); N. falcata: 19 (x 1250); N. hirsutula: 20 (x 1250), 21 (x 1000); N. lauterbachii: 
22 (x 1250); N. pectinata: 23 (x 1000); N. radicans: 24 (x 1250). 


WSS 
SSS 


j 
j 


Plate III. Spore characteristics of Oleandra. 


Figs. 25: Group of spores within a broken sporangium: 26: typical structure of a spore 
with broken perine; 27: a partly spinulose and partly ridged spore: 28: a wholly spinulose 
spore; 29: another type of spinulose spore with thick ridges; 30, 3/, 32: close-up views of 
some broken perines, showing erect blunt rods and details of surrounding structures: 33, 34 
close-up views of some unbroken perines. showing holes and short spines; 35: structur 
the short spines, etc. on the perine. 

O. undulata: 25 (x 150), 26 (x 1000), 29 (x 500), 37 (x 1250), 32 (x 4000), 35 
O. wallichii: 27 (x 1000), 28 (x 750); O. madagascarica: 33 (x 500); O. pistillaris: 30 ( 
34 (x 500). 


re oO 


43 44 45 


Plate IV. Some representative spores of Oleandra species. 


O. Madagascarica: figs. 36 (x 500), 37 (x 1250); O. neriiformis: 38 (x 1000), 39 (x 1500), 
40 (x 1000); O. pistillaris: 41 (x 500); O. undulata: 42 (x 1000); O. wallichii: 43 (x 1000), 
44 (x 1000), 45 (x 750). 


Plate V. Spore characteristics of Arthropteris. 


Figs. 46: Group of spores seen under low magnification; 47, 48: normal spores and 
part of an ablated spore; 49, 50, 5/, 52: close-up views of the spores, showing detailed 
structures of various kinds of outgrowths on the perines; 53, 54: broken spores, showing 
rough perines and smooth exines. 


A. articulata: figs. 46 (x 250), 47 (x 750), 48 (x 750), 49 (x 2500); A. pallisotit: 50 (x 2500), 
53 (x 750); A. tenella: 51 (x 2500); A. beckleri: 52 (x 1250); A. dolichopoda: 54 (x 1250) 


Plate VI. Some representative spores of Arthropteris species. 
A. beckleri: figs. 55 (x 500), 56 (x 500); A. dolichopoda: 57 (x 500). 58 (x 500); A. 


monocarpa: 59 (x 250); A. obliterata: 60 (x 750); A. pallisotii: 61 (x 750), 62 (x 500); A. 
repens: 63 (x 500); A. tenella: 64 (x 1000); A. wollastonii: 65 (x 500), 66 (x 500). 


SAURAUIAE GERONTOGEAE 


I. Notes on Malayan species 
by 


R. D. HOOGLAND 
Australian National University 


Canberra 


Abstract 


Saurauia tristyla has been erroneously recorded from Malaya; the correct placement of 
these records is discussed. §. roxburghii Wall. and S. napaulensis DC. are new records for 
the area. S. pentapetala (Jack) Hoogl. is proposed as a new combination, with basionym 
Ternstroemia pentapetala Jack. Three new species are described: S. fragrans Hoogl., S. 
mahmudii Hoogl., and S. malayana Hoogl. S. cauliflora var. calycina King is placed in 
synonymy under S. leprosa Korth., and S. grandis Ridl. in synonymy under S. vulcani Korth. 
The altitudinal distribution of the 10 species found in Malaya is discussed. 


Introduction 


Under the general title SAURAUIAE GERONTOGEAE the author hopes 
to present a number of papers on the Old World species of Saurauia. The present 
must be seen only as one which is preliminary to a fully documented account of 
the Malayan species to be presented in the near future. Other accounts currently 
being prepared cover the species from continental South-East Asia, Java, and the 
Lesser Sunda Islands. Considering the high degree of local endemism in the genus, 
the preparation of separate treatments for the major islands or island groups of 
Malesia offers the opportunity to gradually compound our knowledge of the genus 
in this region. 

The author is grateful to the authorities of the following herbaria for their 
ala in allowing the study of their collections during visits or by way of 
oans: 


Botanical Section, Department of Agriculture, Bangkok (BK). 

Forest Herbarium, Royal Forest Department, Bangkok (BKF). 

British Museum (Natural History), London (BM). 

Herbarium Bogoriense, Bogor (BO). 

peor ecamem Herbarium, Botanical Survey of India, Sibpore, Howrah 

Herbarium Australiense, C.S.I.R.O., Canberra (CANB). 

Royal Botanic Garden, Edinburgh (E). 

Conservatoire et Jardin botaniques, Genéve (G). 

Royal Botanic Gardens, Kew (K). 

Forest Research Institute, Kepong (KEP). 

Department of Botany, University of Malaya, Kuala Lumpur (KLU). 

Rijksherbarium, Leiden (L). 

Laboratoire de Phanérogamie, Muséum National d’Histoire Naturelle. 
Paris (P). 

Botanic Gardens, Singapore (SING). 

De ore we Universiti Kebangsaan Malaysia, Kuala Lumpur 


111 


112 Gardens’ Bulletin, Singapore — XXX (1977) 


Fieldwork in Malaya was made possible with generous assistance from the 
University of Malaya, Kuala Lumpur; the Forest Research Institute, Kepong; the 
Universiti Kebangsaan Malaysia, Kuala Lumpur; and the Botanic Gardens, Penang. 


I am indebted to Mrs. Cathy Porter for the illustrations of Saurauia mahmudii 
and §. malayana, and to Miss Jill Ruse for those of S. fragrans. 


I. Saurauia tristyla auct. non DC. in Malaya 


The type of Saurauia tristyla DC, [in Mem. Soc. Phys, Hist. Nat. Genéve 1 
(1822) 423, t. vii] originated from the Moluccas and represents a species which 
has subsequently been collected repeatedly there and in Celebes (Sulawesi). It has 
also been recorded from a wide geographical area including West New Guinea, 
Sumatra, Borneo, Malaya, Burma, Thailand, Indo-China, and Southern China 
(including Taiwan). Most, if not all, of these further records must be discarded 
as being referable to other species. The present note concerns itself only with the 
records and collections from Malaya and Peninsular Thailand. The main literature, 
under this name and for this area, is contained in King [in J. Asiat. Soc. Beng. 
59 II (1890) 197], Ridley [Fl..Mal. Pen. 1 (1922) 207], Craib [Fl]. Siam. Enum. 
1 (1925) 129], and Keng [in Fl. Thail. 2 (1972) 110]. 


A study of the material in CAL, seen and annotated by King, revealed that 
he included specimens of three species for which I accept the names Saurauia 
fragrans Hoogl., S. pentapetala (Jack) Hoogl., and S. roxburghii Wall. In addition, 
two specimens which I refer to §. pentapetala were annotated S. cauliflora DC. 
(Scortechini 99) and S. pendula Bl. (Scortechini 331) whereas one collection 
(Scortechini s.n.: CAL 49323/4) of S. roxburghii was annotated “quite different 
from S. tristyla DC., G. King’. None of these appear to have been considered 
further in King’s account of the genus in the Materials for a Flora of the Malayan 
Peninsula. The same three species were available to Ridley in the herbaria of 
Singapore and Kew and included by him in S. tristyla, but as S. fragrans has not 
been found in Thailand the accounts of S. tristyla by Craib and Keng include only 
the other two species. 


1. Saurauia pentapetala (Jack) Hoogl., comb. nov. 


Basionym: Ternstroemia pentapetala Jack in Mal. Misc. 1 (5) (1821) 40; cf. 
Merr. in J. Arn, Arb. 33 (1952) 247. = Cleyera pentapetala (Jack) Spreng., Linn. 
Syst. Veg. (ed. 16) 2 (1825) 596. = Scapha pinangiana Choisy in Mém. Soc. Phys. 
Hist. Nat. Genéve 14 (1855) 119, nom. illeg. — TYPE: ‘“‘Native of Pulo Penang”’, 
lost; NEOTYPE: Hoogland 12601, SE. slope of Penang Hill, Penang, 8 April 
1975, on forest edge at ca 420 m altitude, in CANB, duplicates being distributed 
to K, KEP, L, SING, US. 


Synonym: Saurauia nudiflora var. angustifolia Craib, Fl. Siam. Enum, 1 (1925) 


128. — TYPE: ‘“PATTANI. Bachaw, c. 600 m., by stream in evergreen forest, 
Kerr, 7196”; HOLOTYPE in ABD, n.v., ISOTYPES in BK, BM, E, K. 


Misapplied name: Saurauia tristyla auct. non DC.: Wall., Numer. List (1829) 
no. 1466 et Pl. Asiat. Rar. 2 (1831) 40; Dyer in Fl. Br. Ind. 1 (1874) 287, p.p.; 
King in J. Asiat. Soc. Beng. 59 II (1890) 197, p.p.; Ridl., Fl. Mal. Pen. 1 (1922) 
207, p.p.; Craib, Fl. Siam. Enum. 1 (1925) 129, p.p.; Burk. & Henders., Gard. 
Bull. Straits Settl. 5 (1925) 349, p.p.; Henders., ibid. 4 (1928) 226, p.p.; Burk., 
Dict. Econ. Prod. Mal. Pen. (1935) 1967 et ed. 2 (1966) 2002, p.p.; Keng in FI. 
Thail. 2 (1972) 110, p.p. 


The two earliest names, validly published for Old World species of Saurauia, 
are those given by W. Jack under the generic name Ternstroemia to one Sumatran 
and one Malayan species.’ The Sumatran species was correctly renamed Saurauia 
rubiginosa (Jack) Merr., but the required new combination has never been proposed 


Malayan Saurauiae 113 


for the Malayan species. The first author to have placed it in Saurauia appears to 
have been Dyer (1874) who lists it in synonymy under S. tristyla DC. This species 
had already been recorded from Penang, the type area for Jack’s name, by Wallich 
(1829, 1831) on tke basis of a specimen collected by Porter. All subsequent 
authors followed this placement of the name. Merrill, in his account of William 
Jack’s genera and species of Malayan plants (1952), did not specifically place it 
beyond saying that it “belongs clearly in the group with S. tristyla DC.” 


Saurauia pentupetala is probably the most common and most widely distri- 
buted species in Malaya, having been found in most of the mainland states and 
on Penang and Singapore islands. In Peninsular Thailand it is known from Ranong 
and Surat Thani districts southward. The species is most commonly found in the 
lowlands, up to ca 500 m altitude, with an occasional occurrence at higher altitudes 
(up to 1100 m). 


2. Saurauia roxburghii Wall., Numer. List. (1829) no. 1467, nom. nud., et PI. 
Asiat. Rar. 2 (1831) 40. — TYPE from India. 


Misapplied name: Saurauia tristyla auct. non. DC.: King in J. Asiat. Soc. Beng. 
59 II (1890), 197, p.p.; Ridl. in Trans. Linn, Soc. London II Bot. 3 (1893) 277, 
et Fl. Mal. Pen. 1 (1922) 207, p.p.; Craib, Fl. Siam. Enum. 1 (1925) 129, p.p.; 
Burk. & Henders., Gard. Bull. Straits Settl. 3 (1925) 349, p.p.; Henders., ibid. 4 
(1928) 226, p.p.; Burk., Dict. Econ. Prod. Mal. Pen. (1935) 1967 et ed. 2 (1966) 
2002, p.p.; Keng in Fl. Thail. 2 (1972) 110, p.p. 


Saurauia roxburghii is a widespread species of continental South-East Asia, 
extending from eastern Nepal through southern China to the Ryukyu Islands, 
through Burma, Thailand, and Indo-China, and is now for the first time recorded 
for Malaya. Here it is less widespread than S. pentapetala, being absent from 
Malacca, Johore, Singapore, and Penang, and, though it has been found at quite 
low altitudes, tends to be more frequent between 500 and 1000 m where it may be 
common in regrowih vegetation; the maximum recorded altitude is 1250 m. 


3. Saurauia fragrans Hoogl., sp. nov. (fig. 2) 


Arbor parva usque 12 m alta, 20 cm diametro. Folia oblonga vel obovato- 
oblonga, plerumque 18-30 cm longa, 6-11 cm lata, 14-17-nervata, apice acuta + 
acuminata, basi acuta vel obtusa, petiolo 24 cm longo. Flores solitarii vel usque 
20 fasciculati, interdum inflorescentia cymosa paucifiora interspersa, in axillis 
foliorum vel cicatricium foliorum. Pedunculus usque 5 mm _ longus, pedicellus 
§-22 mm longus. Sepala suborbicularia, 3-5 mm longa et lata. Corolla late 
campanulata (fere plana), 2 mm.alta, 13 mm diametro; tubus 14-2 mm longus; 
lobi 51-84 mm longi, 44-74 mm lati, apice bilobulati. Stamina plerumque 20-23, 
filamentis subaequalibus 23-34 mm longis, antheris versatilibus dorsifixis 1}-2 mm 
longis. Ovarium subglobosum, 1.8-2.3 mm altum, 1.8-2.7 mm diametro, dense 
sericeo-hirsutum, triloculare; styli 3-44 mm longi, basi } mm connatl. 

Squamae triangulares usque 0.7 mm longae, 0.3 mm latae, apice acuminatae, 
superficie dense trichomatosae, in ramellis, pendunculis, pedicellis, et foliis, 
praesertim facie inferiori; inter squamas pili crassiusculi usque 0.3 mm_ longi, 
frequenter stellatim collocati. 


TYPE: Hoogland 12571, along Kuala Lumpur—Raub road between m.s. 53 and 
54 (ie. near The Gap), Selangor, 27 March 1975, in fairly tall secondary forest 
on roadside at ca 720 m altitude. HOLOTYPE in CANB, ISOTYPES being 
distributed to A, BO, E, G, K, KEP, L, MEL, SING, US. 

Further collections examined: Whitmore FRI 2622 (L. SING), Kasim & 
Rahim K 750 (UKM), Kunstler 10638 (CAL, K), Teo & Pachiapa 340 = KL 
2904 (K, L, SING), Lye L 68/71 (SING), Poore 856 (KLU), Burkill SF 3184 
(SING), Stone 6894 (KLU). 


114 Gardens’ Bulletin, Singapore — XXX (1977) 


Misapplied name: Saurauia tristyla auct. 
non DC.: King in J. Asiat Soc. Beng. 59 
II (1890) 197, p.p.: Ridl., Fl. Mal. Pen. 1 
(1922) 207, p.p.; Burk., Dict. Econ. Prod. 
Mal. Pen. (1935) 1967 et ed. 2 (1966) 
2002, p.p. 


Saurauia fragrans is known only from 
a few localities in Selangor: The Gap 
(straddling the Selangor—Pahang bound- 
ary), Kepong, Ulu Bubong, and Ulu 
Gombak, and one in Negri Sembilan: 
Gunong Tampir, occurring at elevations 
from 75 to 750 m. Outside Malaya it has 
also been found in Northern Sumatra. 


The specific epithet has been chosen 
as, in contrast with other . species of 
Saurauia encountered by me in India, 
Malaya, Java, Flores and New Guinea, 
the flowers of the type specimen were 
heavily sweet-scented, a feature made 
even more noticeable because of the very 
large number of flowers present on the 
tree. The fragrance of the flowers was 
also noted in Lye L 68/71. 


The differences between Saurauia 
roxburghii, S. pentapetala, and S. fragans 
are set out in the accompanying table. 


S. fragrans 
1 (-3)-flowered, with at most short peduncle; 


at least those below the leaves in cluster of up 


to 20. 
trichomes on base of and between scales. 


S. pentapetala 


INFLORESCENCE 
1-3 (-5)-flowered, in most plants predominantly 
PEDUNCLE AND PEDICELS 


1-flowered, with at most short peduncle; at least 


those below the leaves in cluster of up to 15. 
incorporated in base of scales and sparse or 


mostly absent between scales. 


S. roxburghii 
With tomentum of very short villose hairs and | With broad-based scales, with villose tomentum | With long fairly broad based scales and short 


peduncle; at least those below the leaves in 


(1-) 3-7-flowered, often open with relatively long 
cluster of up to 4. 


occasionally a few small elongated scales. 


=—_— 


115 


‘umouyun Areao Buningy oinyepy 
‘1199 19d Q9Z-OpZ BO JOquINU 9[NAGD 


‘Iomoy usdo ut 
duipvoids Apyysis ‘tomoy uodo Ajsvou ur yydresys 
puv yo19 07 pnq Ajsivo Ul yoOWUOD sue 9[A\g 


‘du[noo|-(p-) € (-Z) 
(duo, wut ¢() 07 dn saivy) oynsary Ajosuod 


‘xodu 78 pYyiq ‘O[NVS19A ‘poxyissop ss9yjUuy 
‘snoigels syusUEyi 
9C°8 1 


‘png Ul UOTPATSOR 410]U09 JO SALMR 
‘xodu jo o[ppiw ul UOIsIOUL MOT[eYs YIM soqgo’y 
‘(WILL ()[-Z BO SL YOIYM) vI[O109 JO YBud] [v}0) Jo 
sso] 10 ¥ ynoqe Ajuo ‘oye[nuUeduIEd-proiqg oqny 


‘umouyun xA[vo Bunini.y 


‘OPISINO 9SOJUDWIO}-97e[ [918 
Ajouy png ul posodxs o194M yseo, ye ‘suo, Ww 
G-€ vO ‘YO, Ul Av[IUNIs sjedos JouUL puv 19jnNO 


Malayan Saurauiae 


1") wu FL 
vo Aivao Bunning sine 


‘1199 19d OSE-OOE BO JoquinuU dINAG 
‘Iomoy usdo ut Juipvoids 
usdo Ajivou ut yydiesys 

[4v9 Ul JOWUOD sULIe 9[AISg 

‘aB[Ndo[-($-) p-€ 
‘$NOIQL[H 


vo “ueIp wu [| 


Apyays Ajuo ‘1aMo 
pue yoo19 OF png 


‘xode Jv pylq ‘a[NesI9DA ‘poxyisiop si9yjUYy 
‘snoiges syuoWeyi.y 
vY-8C 


SNAWVLS 


‘png UL UONLATSOR 110}U09 Jo shemie 
‘xode JO d[ppiul Ul UOISIOUL MOT[VYS YIM sogo7y 
‘(WIL ¢[-8 VO SI YOIYM) VI[OLOD Jo YWuUdT [v}0} JO 
sso] JO — ynoge AjuoO ‘oJe[NURdUIeO-prOoIg 9qny] 


v1Ti010Dd 


“A1GAO 
duninsgy wos Avme poainos Ajjensn ysed soddn 
ui ‘ArvaAo Buns 0} YIBUd, Ul JeprUNs “Indy UT 
OZIS [VUISIIO SOLU) Z-F[ 0} podIejus AppouNsiG 


‘opisyno snoiqge[s ‘duo, WU g-¢ 
vo ‘yydus, Ul sejiwis sjedos Jouut pur s9jnO 


S1vdd§ 


11%) pue ‘weip wut g ed AivAo Buyninsy singe 
‘1199 sod QZ[-0R BO JoquInU a;nNAG 


‘IaMOy Uusdo UI OF OBR} 
pnq Ajivo wos Aljeyuozis0y Buipvoids sue 9[Ajg 


‘4RND0]-¢-£ 
‘snoiqr|y 


‘xode 7e O8njo1 jsow 
ye ‘Q[VRSIZA JOU ‘poxyiseq sloyUY “ed JOMOT 
ur Ajurur ‘siiey BuO, [eI9ANS YUM syuoWwelLy, 


Ly-ve 


A A A 


‘Wo UOD A[oivs ‘UOTRAISOR 
sutAivaA JO png ut ‘xode ye s[pprum ul Uorsiour 
MO[[eYys YM Ajoie1 ‘popunos Ajjensn soqgo7y 

‘(wu 1-9 BO SI YoIymM) 
B[[O109 JO YSUd| [v}0} JO | ynOg”E ‘a}e[OOOIN 9qny 


‘ALVAO Buyinsy 9INyeu UY} JoyoYs ApjouN 

-sIp pue 0} possoidde ‘yinsy ul podivpus Ajpieyy 
“opis}no 

WINJUDWIO} DSO]IIA JIOYS 9WOS YIM png Ul posodxe 
DIDYM yseo] ye “Buol ww 7'¢-g'¢ ‘dsol ('p-9'Z vO 
‘souo sJouul Jo ¢/Z ynoge YsUs] UI souUO J9jNO 


116 Gardens’ Bulletin, Singapore — XXX (1977) 


II. Saurauia in the Cameron Highlands 


During a short visit to the Cameron Highlands in May 1973 I was able to 
collect a number of specimens of Saurauia. On closer examination it became clear 
that amongst these collections two undescribed species were represented. When IL 
tevisited the area in March-April 1975, I took the opportunity of collecting further 
specimens of these two species and making notes of their altitudinal occurrence. 
Similarly, the altitudinal zonation of the other species in the area and along the access 
road from Tapah to Tanah Rata was noted and in part documented by the collect- 
ing of herbarium specimens. It appeared that out of ten species now known to 
occur in Malaya, eight are found in the area covered. Only S. rubens Ridl. and 
S. fragrans Hoogl., both of limited known occurrence, are absent. 


altitude in metres 


o0osl 


Ss 
e) 
°o 


009 


fragrans 

leprosa 

mahmudii 
Fig. 1. Known alti- 

malayana tudinal range of 
Saurauia species in 
the Cameron High- 

napaulensis lands (solid block) 
and elsewhere in Ma- 
laya (open block). 

nudiflora 

pentapetala 

roxburghii 

rubens 

vulcani 


In fig. 1 the observed altitudinal range in the Cameron Highlands and along 
the road from Tapah is presented along the altitudinal range outside the Cameron 
Highlands of all species known from Malaya as ascertained from data available of 
herbarium specimens. The’ species are arranged in alphabetical order; apart from 
those already treated the following are involved: 


ie Ah oo - 


_ - 


117 


Malayan Saurauiae 


; b. and c. open flower, x 2; 


2 
d. flower bud just before opening, x 3; e. longitudinal section of flower, x 2; f. stamens, 


nm 


Saurauia fragrans Hoogl. a. flowering branch, x 


Fig. 2. 


x 4, All after Hoogland 12571. 


118 Gardens’ Bulletin, Singapore — XXX (1977) 


4. Saurauia leprosa Korth., Verh. Nat. Gesch. Kruidk. (1842) 131. — TYPE from 
Java. 


Synonym: Saurauia cauliflora var. calycina King in J, Asiat. Soc. Beng. 59 II 
(1890) 199; Ridl., Fl. Mal. Pen. 1 (1922) 208; Craib, Fl. Siam. Enum, 1 (1925) 
127; Henders. in J. Mal. Br. Roy. Asiat. Soc. 17 (1939) 25, 37; Keng in Fl, Thail. 
2 (1972) 110. — TYPE: ‘‘Perak: Batu Kurau. Scortechini, No. 1614.”; HOLO- 
TYPE in CAL, ISOTYPE in SING. 


Misapplied name: Saurauia cauliflora auct. non DC.: Burk. & Henders, in Gard. 
Bull. Straits Settl. 3 (1925) 336, 349; Henders. ibid. 4 (1928) 226; Burk., Dict. 
Econ. Prod. Mal. Pen. (1935) 1967 et ed. 2 (1966) 2002. 


This is in Malaya a rather rare mainly lowland species found only in Perak, 
Pahang, and Selangor, and occurs outside Malaya in Peninsular Thailand, Sumatra, 
and Java. Throughout its range some individuals are almost exclusively cauliflorous, 
others are exclusively ramiflorous or carry the inflorescences in the leaf axils, while 
few individuals show the whole range of inflorescence positions. 


5. Saurauia mahmudii Hoogl., sp. nov. (fig. 3) 


Arbor parva usque 6 m alta. Folia obovata vel obovato-oblonga, plerumque 
20-30 cm longa, 10-15 cm lata, 20-28-nervata, apice obtusa late acuminata, basi 
acuta subdecurrentia, petiolo 24-44 cm longo. Inflorescentiae solitariae vel 2-4 
fasciculatae in axillis foliorum vel cicatricium foliorum, pro parte majori dichasia 
triflora, nonullae biflorae vel uniflorae, usque 7 cm longae. Pedicellus 2-5 cm 
longus. Sepala 9-12 mm longa, 8-10 mm lata, basi 3 mm crassa. Corolla late 
urceolata, 13-15 mm longa, 15 mm diametro; tubus 7-9 mm longus, 12 mm 
diametro; lobi 8-10 mm longi, 74-11 mm lati, recurvati, apice bilobulati. Stamina 
plerumque 25-32, filamentis subaequalibus 5-53 mm longis, antheris versatilibus 
dorsifixis 24-3 mm longis. Ovarium semi-globosum 3-34 mm altum, 5 mm diametro, 
glabrum, triloculare; styli 7 mm longi, basi 14-2 mm connati. 


Squamae anguste triangulares, 0.5-1.0 mm longae, 0.2-0.3 mm latae, acutae, 
margine praesertim) inferiori piloso. In ramis, foliis, pedunculis, pedicellis, bracteis, 
et bracteolis sparsae; in sepalis absunt, interdum uno vel duabus basi exceptis. 


TYPE: Hoogland 12579, along road to summit of Gunong Brinchang near m:s. 
46. Cameron Highlands, Pahang, 1 April 1975, in gully on steep slope at ca 1770m 
altitude. HOLOTYPE in CANB, ISOTYPES being distributed to BRI, K, KLU, 
L, SING, US. 


Further collections examined: Whitmore FRI 15449 (KEP), Hoogland 12326 
(CANB, K, KEP, L), 12327 (CANB, SING), /2328 (CANB, SING). 12580 
(CANB. L), 12581 (A, BO, CANB, KEP), Kasim & Zainudin K 988 (UKM), 
Stone 5595 (KLU); all from the Gunong Brinchang area. 


The species closely resembles §. nudiflora from which it is readily distinguished 
by the glabrous ovary as against densely sericeous. In S. nudiflora nearly all flowers 
are solitary though a few 2- or 3-flowered inflorescences occur in most plants; in 
S. mahmudii the inflorescences are mostly 3-flowered with fewer 2- or 1-flowered 
ones interspersed. Individuals with predominantly 3-locular ovaries are found in 
S. nudiflora, though only one such collection is known to me (Hoogland 12570) in 
which the flowers are smaller, with a smaller than usual number of stamens for this - 
species. In all other collections examined the ovary is predominantly 5-celled, with 
occasional flowers with 4- or 6-celled ovary. Var. tetragyna Boerl., originating 
in Java, is reported to have a 4-celled ovary. In all collections of S. mahmudii the 
ovary is 3-celled (one 2-celled ovary was found in 130 flowers analysed). 


I have dedicated this species to Mahmud bin Sider, of the University of 
Malaya, who assisted me in collecting in the Cameron Highlands in 1973 and 1975. 


Malayan Saurauiae 119 


*, 


a f VE, a 
Camp ?y 
iy: se Oa ‘ 
=4 AY, Dy we tae c = 


DAG 4 


PY 
i¢ 
i 


Fig. 3. Saurauia mahmudii Hoogl. a. flowering branch, x 4; 5. open flower, x 2; 
os Fs eerseeg section of flower, x 2; d. stamens, x 2; e. fruit, x 2. All after Hoogland 


120 Gardens’ Bulletin, Singapore — XXX (1977) 


6. Saurauia malayana Hoogl., sp. nov. (fig. 4) 


Frutex vel arbor parva usque 12 m alta, 20 cm diametro. Folia obovato- 
oblonga, plerumque 14-32 cm longa, 6-15 cm lata, 12-20-nervata, apice obtusa 
breviter acuminata, basi rotundata vel obtusa, petiolo 14-44 cm longo. Inflores- 
centiae solitariae ve] 2-3 fasciculatae in axillis foliorum vel cicatricium foliorum, 
dichasiales, 3—20-florae, usque ad 20 cm longae. Pedunculus 1-10 cm longus. 
Pedicellus 5-40 mm longus. Sepala 2 exteriora late triangulari-ovata, 8-9 mm 
longa, 9-12 mm lata; 2 interiora suborbicularia, 8-10 mm longa, 9-12 mm lata; basi 
2 mm crassa. Corolla campanulata, 10-15 mm longa, 16-20 mm diametro; tubus 
5-7 mm longus, 10-13 mm diametro; lobi 84-11 mm longi, 9-13 mm lati, 
aliquantum reflexi, apice bilobulati. Stamina plerumque 30-40, filamentis 
subaequalibus 34-54 mm longis, antheris versatilibus dorsifixis 2-3 mm _longis. 
Ovarium semi-globosum 24-34 mm altum, 44-6 mm diametro, dense hirsutum, 
(4-) 5 (-7)-loculare; styli 54-74 mm longi, basi 1-24 mm connati. 

Squamae anguste triangulares acuminatae, superficie trichomatosae; inter 
squamas pili gracilcs stellatim collocati. Squamae ramorum usque 4 mm longae, 
i mm latae (plerumque minores), foliorum usque 3 mm longae, pedunculorum et 
pedicellorum usque 14 mm longae, faciei exterioris sepalorum exteriorium usque 
1 mm longae. Pili graciles usque 1 mm longi. 


TYPE: Hoogland 12591, along road from Ringlet to Tanah Rata between m.s. 
29 and 30, Cameron Highlands, Pahang, 2 April 1975, in open secondary forest in 
gully on steep slope at ca 1150 m altitude. HOLOTYPE in CANB, ISOTYPES 
being distributed to A, G, K, KEP, L, SING, US. 


Further collections examined: Hoogland 12332 (CANB), 12335 (CANB, 
KLU, L), 12576 (A, BO, CANB, E. KEP), 1/2586 (BO, CANB, E, KLU, L, 
US), 12587 (CANB, L), 12588 (CANB, L), 12590 (CANB, G, K, SING), 12595 
(BH, E, CANB, KLU), Turnau 789 (KLU). 


Amongst its Malayan congeners this species stands out by the rather long, 
open inflorescences with large flowers. Other species in the area with many- 
flowered inflorescences are S. napaulensis with much smaller flowers and S. vulcani 
with the flowers in a much more compact inflorescence. Both these species have 
generally many more flowers to each inflorescence, and also differ substantially in 
the nature of their scales, 


Saurauia malayana has been found only in the Cameron Highlands and is 
particularly common around the Ringlet between 1100 and 1300 m altitude, 
whereas a few trees were also found near Tanah Rata at 1400 m. 


7. Saurauia napaulensis DC. in Mém. Soc. Phys. Hist. Nat. Genéve 1 (1822) 421. 
— TYPE from Nepal. 


This is another widespread species of continental South-East Asia, extending 
from north-western India (Himachal Pradesh) to southern China, through Burma, 
Thailand, and Indo-China, and now, like S$. roxburghii, recorded for the first time 
for Malaya. Compared to that species, it ranges further West in the foothills of the 
Himalayas, but not as far East in the Chinese region. While I have no hesitation in 
accepting S. roxburghii as indigenous in Malaya, there must be some reservations 


about this for S. napaulensis. The species was not collected in the Cameron . 


Highlands (the only area in Malaya where it has been found until 1936) but it is 
now extremely common, particularly in disturbed areas between Tanah Rata and 
Brinchang and on the summit of Gunong Brinchang. 


The species is grown as an ornamental in the tea-growing areas of Ceylon and, 


in particular, of Southern India and the possibility of its introduction into the 
Cameron Highlands for the same purpose cannot be discarded. On the other hand, 


Malayan Saurauiae 121 


| 
: 

<< 
4 


Fig. 4. Sauranuia malayana Hoogl. a. flowering branch, x 4; 6. open flower, x y 
c. longitudinal section of flower, x 2; d. stamens, x 2; e. imature fruit, x 2. All after 
Hoogland 12591. 


122 Gardens’ Bulletin, Singapore — XXX (1977) 


it may have occurred there naturally and extended its area substantially with the 
opening up of the highland country, The nearest known natural occurrences 
outside Malaya are in Thailand on Doi Inthanon in Chieng Mai district and Phu 
Miang in Phitsanulok district, some 1500 km to the North. 


8. Saurauia nudiflora DC. in Mém. Soc. Phys. Hist. Nat. Genéve 1 (1822) 422, 
t. v; King in J. Asiat. Soc. Beng. 59 II (1890) 198; Ridl. in Trans. Linn. Soc. 
London II Bot. 3 (1893) 277, in J. Fed. Mal. St. Mus. 4 (1909) 6, et Fl. Mal. 
Pen. 1 (1922) 207; Burk. & Henders. in Gard. Bull. Straits Settl. 3 (1925) 336, 349; 
Henders., ibid. 4 (1928) 226; Burk., Dict. Econ. Prod. Mal. Pen. (1935) 1967 et 
ed. 2 (1966) 2002. — TYPE from Java. 


Common in north-western Malaya, where it has been found in Kedah, Perak, 
Pahang, and Selangor. Outside Malaya it occurs in Sumatra and Java. 


9. Saurauia rubens Ridl. in J. Roy. Asiat. Soc. Straits Br. 61 (1912) 1, et FI. 
Mal. Pen. 1 (1922) 208. — TYPE: “Selangor: Semangkok Pass, at the foot of 
the hill leading to the Sempang mines.”; HOLOTYPE in SING (Ridley 15771, 
‘““Sempang track, Ap. 1911’’), ISOTYPE in BM. 


Common in the Fraser’s Hill area, also collected once in Trengganu 
(Mandi Angin). Endemic to Malaya. 


10. Saurauia vulcani Korth., Verh. Nat. Gesch. Kruidk. (1842) 128. — TYPE 
from Sumatra. 


Synonym: Saurauia grandis Ridl. in J. Fed. Mal. St. Mus. 4 (1909) 6 et Fl. Mal. 
Pen. 1 (1922) 207. — TYPE: ‘“‘Tel6m River banks.”; HOLOTYPE in SING 
(Ridley 13580, ‘““‘Telom, Nov. 1908”), ISOTYPES in BM, K. 


Apart from its abundant occurrence in the Cameron Highlands, this species 
is known from Malaya only from a recent collection from the Genting Highlands. 
It spreads readily on disturbed roadsides and on landslides. 


The Cyperaceae Tribe Cypereae of Ceylon 


TETSuO0 KOYAMA 
The New York Botanical Garden, Bronx, New York 


Abstract 


This taxonomic treatment, revising all the Ceylonese taxa of the Cyperaceae tribe 
Cypereae, enumerates 59 species of four genera, Cyperus, Pycreus, Mariscus and Kyllinga. The 
following new names are proposed: Cyperus compressus ssp. micranthus, C. diffusus ssp. 
macrostachyus, C. nutans ssp. eleusinoides, Pycreus flavidus, P. pumilus ssp. membranaceus, 
Mariscus cyperinus ssp. laxatus, M. pedunculatus, M,. sumatrensis, and Kyllinga odorata 
ssp. cylindrica. 


In the course of my preparation of a taxonomic account of the Cyperaceae for 
the Flora of Ceylon, a project under the auspices of the Smithsonian Institution 
(Koyama 1970), I have had the opportunity of revising important historical 
collections of Ceylonese Cyperaceae that are housed in the Herbarium of the 
National Botanic Gardens, Peradeniya. These old collections, of which only a part 
is represented at Kew and elsewhere, have not received the attention of specialists 
since Trimen published his Handbook (1900). In addition, a series of field expedi- 
tions recently sent to Ceylon (Koyama 1970, 1974) resulted in an accumulation 
of a substantial number of critical gatherings, among which are several cyperoid 
taxa that are new to the flora of Ceylon. In the present paper 59 species of the 
tribe Cypereae are recognized as valid for the flora of Ceylon and are classified 
under four genera, Cyperus, Pycreus, Mariscus and Kyllinga. The treatment not 
only identifies the collections mentioned above but also updates the nomenclature 
with particular emphasis on the coordination to the two recent imporant works on 
the Cypereae (Kiikenthal 1935-36; Kern 1974). The remarkable richness of 
cyperoid taxa in Ceylon rests on the fact that besides the so-called Indo-Malesian 
and the Pantropic elements, on this little island there occur Western Asiatic elements 
reaching Ceylon as well as those interesting endemic ones which are confined to 
the southern part of Deccan Peninsula and Ceylon. Special attention was paid to 
these entities of phytogeographic interest. 


I appreciate the assistance of Dr. F.. Raymond Fosberg, who has given me 
this opportunity of studying Ceylonese Cyperaceae. Messrs R. Cooray, T. Herat, 
N. Balakrishnan, A. H. M. Jayasuria and K. Sumithraarachchi certainly deserve 
my sincere thanks for their help in the field. The financial assistance from the 
Smithsonian Office of International Program is gratefully acknowledged for their 
research grant in defraying my field expenses. 


KEY TO GENERA 


1. Rhachilla not articulated, persistent, hence glumes falling apart from rhachilla 
2. Achenes triquetrous or dorsi-ventrally flattened with one side facing 


PEE ae eR WE this dw cn in Res Viwddiey tn onan- ss ess. i ARNE TO fo Cyperus 
2. Achenes laterally flattened with one angle facing rhachilla ......... Pycreus 
1. Rhachilla articulated, hence spikelets falling in entirety 
3. Achenes triquetrous with one side facing rhachilla .................. Mariscus 
3. Achenes laterally flattened with one angle facing rhachilla ......... Kyllinga 


123 


124 Gardens’ Bulletin, Singapore — XXX (1977) 
CYPERUS L. 


1. Stigmas 3, or style more or less undivided (sp, no. 33) 
2. Spikelets spicately disposed on conspicuously elongated rhachis, thus 
forming spikes 
3. Rhachilla of spikelets winged with base of glumes, which is decurrent 
along the rhachilla internode 


4. Spikelets many to numerous; spikes with long rhachis, cylindrical, 
normally 3-7 cm long 
5. Leaves with elongated blade 
6. Spikelets linear, terete or subterete, 0.5-1 mm wide 
7. Spikelets patent to spreading; spikes as a rule peduncl- 
ed; leaves 4-15 mm wide; culm smooth on angles 
Pe ERC Ta ae EE Penne VET 1 C. digitatus 


7. Spikelets erect to suberect; spikes subfastigiate, sub- 
sessile; leaves 20-30 mm wide; culms scabrous on 


upper,angles 14:03..0-00.n2 zene 2 C. platyphyllus 

6. Spikelets lanceolate to lance-oblong, ca, 1.55 mm wide, 
flattenéd 27.28.2508). QIAN. AR RA 3 C. exaltatus 

5. Leaves reduced to bladeless sheaths ............... 4 C. papyrus 


4. Spikelets several to teens, occasionally up to 30; spikes with rather 
short rhachis broadly ellipsoid or ovoid, normally 1-4 cm long 


8. Plants with creeping rhizomes or stolons (except in C. bulbosus, 
which bears tunicate bulb at base of culm) 

9. Culms thick, normally more than 5 mm wide and 60 cm 
tall, clothed at base with bladeless sheaths 

10. Culms obtusely trigonous to subterete; glumes lance- 

ovate, acute-tipped; bracts much shorter than corymb 

11. Culms not septate ............... 5 C. corymbosus 

11. Culms transversely septate ...... 6 C. articulatus 


10. Culms acutely triquetrous; glumes oblong, obtuse at 
apex; bracts much surpassing corymb ...................+ 


9. Culms slender, mostly 1-2 mm wide and less than 60 cm 
tall; leaves with elongated blade 


12. Culms arising from bulbose base, tufted ............... 

a pod ea eins 04 Wags csborase skp su aaw) eens ann 8 C. bulbosus 

12. Stolons present; culm bases forming a corm-like 
enlargement | 

13. Glumes 2-2.5 mm long; spikelets slightly swollen; 

leaves normally as long as the culm ............... 

« « na'ybamcaiene Qanicle eoniesalt ne tama 9 C. stoloniferus 

13. Glumes 3-4 mm long; spikelets strongly flattened; . 
leaves normally much shorter than the culm 

14. Culms 10-30 cm tall; bracts as long as or 

shorter than the inflorescence ............... 

2D. LEE. ROG 10 C. rotundus 

14. Culms 30-60 cm tall; bracts 2 to 3 times 

as long as the inflorescence 


Ceylon Cypereae 


125 


15. Spikelets linear-lanceolate, gradually 
narrowed to an acute apex, 2.2-3 mm 
wide; glumes ovate 


16. Glumes 3-3.5 mm _ long, red- 
brown, close together not ex- 
posing achene between two 
glumes; spikelets 2.2-2.5 mm 
a 11 C. tuberosus 


16. Glumes 4 mm long, yellow- 
brown, slightly spaced exposing 
portion of achene between two 
glumes; spikelets 3-3.2 mm wide 
CER er AT aes 12 C. retzii 

15. Spikelets linear, rather suddenly 
obtusish at apex, 1.5-2 mm _ wide 
glumes oblong ......... 13 C. scariosus 


8. Plants without creeping rhizomes or stolons 


17. Spikelets linear to lanceolate, 15-2 mm wide; weakly 
flattened with two acute edges; glumes close together, 
acute at apex 


18. 


18. 


Glumes distinctly colored with red-brown, brown or 
dark brown; plants perennial with culms more or 
less enlarged at base 


19. Glumes 2.2-2.5 mm long, pale-green and strong- 
ly colored with dark or purple-brown ............ 
acetate Ae es fade desis n>» <a 14 C. sphacelatus 


19. Glumes 3-3.5 mm long, straw-colored and 
tinged with reddish brown or brown ............ 
OE ny 15 C. tenuiculmis 
Glumes pale or stramineous-green, not at all brown- 


colored; plants annual with fibrous roots only; culms 
not thickened at base .................. 16 C. zollingeri 


Spikelets filiform, subterete, less than 1 mm thick; glumes 
spaced, obtuse at apex 


20. 


20. 


Spikelets spreading, ca. 0.75 mm wide; glumes 
rounded-obtuse at apex with moderately narrow 
ee eee 17 C. distans 
Spikelets erect-patent, ca. 1 mm wide; glumes shallow- 
ly emarginate at apex with very broad hyaline mar- 
(ER Sp Os 18 C. nutans 


3. Rhachilla of spikelets without conspicuous wings, base of glumes not 
decurrent along the rhachilla internode 


Plants perennial with stolons 
22. Rhachis of spikes glabrous; glumes obtuse at white-hyaline 


21. 


21. 


apex 


EE M5, OV cca dwsaccanatacvevans 19 C. procerus 


22. Rhachis of spikes densely hispid; glumes acute at apex 


without broad hyaline apex ......................45 20 C. pilosus 


Plants annual, with fibrous roots only 


126 Gardens’ Bulletin, Singapore — XXX (1977) 


23. Spikelets 1-2.5 cm long, 2-3 mm wide; glumes 2-3.5 mm 
long, acute at apex, tightly disposed 


24. Glumes 3.5 mm long; achenes 1.5 mm long; inflores- 
cence usually open with elongated rays ..................05 
i edienetcLial cicette 21b C. compressus ssp. compressus 


24. Glumes 2 mm long; achenes 1 mm long; inflorescence 
congested in a head-like cluster....0. .::.:2 1:1; «ease 
ey eee 21b C. compressus ssp. micranthus 


23. Spikelets 0.5-1 cm long, 1-1.5 mm wide; glumes 0.75—1.25 
mm long, obtuse at apex, loosely disposed ......... 22 C. iria 


2. Spikelets digitately disposed or capitately congested at apices of inflores- 
cence rays, thus forming heads without rhachis 


25. Inflorescences open with elongated rays 


26. Large-sized perennials with conspicuous rhizome; at least some 
leaves and bracts more than 10 mm wide; culms 30-90 cm tall 


27. Leaves with elongated blade; leafy bracts 5-10 


28. Achenes with acute angles; spikelets in clusters of 2 or 
3, sometimes solitary, weakly compressed 


28a. Spikelets 4-6 mm long, up to 12-flowered ...... 
NitttUaaesieye. cade 23a C. diffusus ssp. diffusus 


28a. Spikelets 10-20 mm long, up to 40-flowered ...... 
qui kh tains 23b C. diffusus ssp. macrostachyus. 


28. Angles of achenes spongy-thickened; spikelets in 
clusters of 5 to 8, strongly compressed .................. 
ae alah « caeiiaclhs 5o. ait ania ie 24 CC.  platystylis 


27. Leaves reduced to bladeless sheaths, leafy bracts more than 
1S: SRR eee 25 C. alternifolius ssp. flabelliformis 


26. Small to medium-sized annuals or short-lived perennials; leaves 
and bracts at most 5 mm wide; culms up to 40 cm tall 


29. Umbel rays longer than or nearly as long as the culm ...... 
LL SLE RS ae FB ND os iacaeteseecene: a an 


29. Umbel rays much shorter than the culm 


30. Glumes awnless; achenes obovate; leaves flattish 


31. Spikelets digitate in clusters of 3-15; glumes 
acutish at apex 


32. Roots brownish; basal sheaths straw-brown 
or pale-brown; spikelets dark green ......... 
een, ee eS 27 C. pulcherrimus 


32. Roots purplish; basal sheaths purple-brown; 
spikelets brown or purple-brown 


Ceylon Cypereae 127 


33. Glumes densely imbricate and close 
together so that achenes not shown 
between glumes; rhizome often con- 
spicuous, rather frequently elongated 
op te a a. Be 28 C. haspan 


33. Glumes sublaxly imbricate and slightly 
spaced, thus achenes exposed between 
glumes; plants tufted with fibrous roots 
Cy ee ee 29 C. tenuispica 


31. Spikelets many, congested in globose heads ...... 
eee. “a EE hs ee Ss eee 30 C. difformis 


30. Glumes awned at apex; achenes oblong; leaves 


canaliculate 
34. Glumes with straight awn ...... 31 C. castaneus 
34. Glumes with recurved awn ...... 32 C. cuspidatus 


25. Inflorescences congested a single head 
35. Style 3-cleft; achenes acute on angles 


36. Rhizome with long creeping stolons; culms 5-20 cm tall; 


hess \1—2 om wide 2.4.2.4 su e..:. 33 C. arenarius 
36. Rhizome short, culms tufted, 20-60 cm tall; heads more 
Watt cml acres “lS... 34 C. conglomeratus 


35. Style scarsely divided at apex; achenes with spongy-thickened 
angles; plants floating on water .................. 35 C. cephalotes 


1. Stigmas 2 


37. Tall perennial; culms ca. 100 cm tall; inflorescences umbelliform with 
elongated rays up to 18 cm long ........................... 36 C. alopecuroides 


1. Cyperus digitatus Roxburgh, [Hort. Beng. 81, 1814, nomen, &] Fl. Indica 
ed. 1, 1: 209, 1820; C. B. Clarke, Fl. Brit. India 6: 618, 1893; Trimen & Hook. 
f., Handb. Fl. Ceylon 5: 36, 1900; Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 55, 
1935; Kern, Fl. Males. I, 7 (3): 601, 1974. 


‘Cyperus venustus R. Br.’ sensu Nees, Contrib. Bot. India 86, 1834. 
‘Cyperus neesii Kunth’ sensu Thwaites, Enum. Pl. Zeyl. 344, 1868. ——- Cyperus 
hookeri Bockeler, Linnaea 36: 308, 1870. ——- Cyperus digitatus Roxb. var. 
hookeri (Béckir.) C. B. Clarke, Fl. Brit. India 6: 618, 1893; Trimen & Hook. f., 
Handb. Fl. Ceylon 5: 37, 1900. 


Jaffna District: Ca. 5 miles W of Paranthan, south shore of Jaffna Lagoon, 
T. Koyama et al. 14043 (NY, PDA, US); Paranthan, Clayton 5263 (K, PDA). 
Anuradhapura District: Wilpattu National Park, East Intermediate Zone, along 


128 Gardens’ Bulletin, Singapore — XXX (1977) 


<< rr ti “‘_OS:;~(‘(‘i~S 


maton Hey _ 


Fig.l. Cyperus platyphyllus Romer & Schultes. AA, habit. B, spikelet with its prophyll. 
C, glume. D, dorsal view of achene. E, ventral view of achene with style. Scales = 1 mm. 


Ceylon Cypereae 129 


the Wildife Dept. trail, ca. 2 miles from barrier gate, T. Koyama & Jayasuria 13965 
(NY, PDA, US). Kandy District: Ramboda, Thwaites C. P. 3043 ex p. (K, PDA): 
Pusselawa, Thwaites C. P. 3043 ex p. (PDA). Colombo District: Ferguson s.n. 
(PDA). 


Distribution. Pantropic with its range extending north into southern China 
and Formosa; abundant in South America and southern Asia. relatively scarce in 
tropical Africa and Australia. 


Plants from India and Ceylon have often been treated as var. hookeri, which 
was segregated by its more copiously spiculose spikes. My observation of this 
species on a world-wide basis does not support this variety. 


2. Cyperus platyphyllus Romer & Schultes, Syst. Veg. 2: 876, 1817: C. B. 
Clarke, Fl. Brit. India 6: 618, 1893, & Illustr. Cyper. ¢. 20, f. 3-4, 1909: Trimen & 
Hook. f., Handb. Fl. Ceylon 5: 38, 1900; C. E. C. Fischer in Gamble, Fl. Madras 
9: 1642, 1931; Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 59, 7. 8&7, 1935. — Fig. 1. 


“Cyperus elatus L.,” sensu Roxburgh, Fl. Indica ed. 1, 1: 207, 1820. 
Cyperus roxburghii Nees in Wight, Contrib. Bot. India 84, 1834; Thwaites, Enum. 
Pl. Zeyl. 70, 1864. Cyperus eminens Klein ex Kunth, Enum. Pl. 2: 70, 
Papyrus latifolia Willdenow. Abhandl. Akad. Berlin 1812/13: 


Trincomalee District: Trincomalee, Glenie in 1864 (PDA). Matara District: 
Kukul Corale, Thwaites C. P. 304] ex p. (K. PDA); Matara, Trimen 23 Feb. 1881 
(PDA). Amparai District: Mile 205 on Rd. A 4,ca. 3 miles N of Pottuvilp, 
T. Koyama et al. 14011 (NY, PDA, US). 


Distribution. Confined to southern India and Ceylon. Marshy places at low 
altitudes. 


This rather rare species is easily discernible by its sessile or nearly sessile 
spikes forming digitate secondary corymbs, and very broad leafy bracts and leaf 
blades. The distribution range of this species, covering the southern part of Indian 
Peninsula and Ceylon only, shows the same pattern as those of Mariscus clarkei, 
Fimbristylis tenuifolia, Carex lobulirostris and Carex leucostachya. 


3. Cyperus exaltatus Retzius, Obs. Bot. 5: 11, 1789: Thwaites, Enum. Pl. Zeyl. 
343, 1864: C. B. Clarke, FI. Brit. India 6: 617, 1893: Trimen & Hook. f.. Handb. 
Fl. Ceylon 5: 32, 1900; Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 64, f. 9 A-F, 
1935; Kern Reinwardtia 2: 99, f. 2, 1952 & Fl. Males. L, 7 (3): 602, 1974. 


Cyperus venustus R. Brown, Prodr. Fl. Nov. Holl. 217, 1810; Thwaites, Enum. 
Pl. Zeyl. 432, 1864. Cyperus altus Nees in Wight, Contrib. Bot. India 84, 
1834. Cyperus exaltatus Retz. var. amoenus C. B. Clarke, Journ. Linn. Soc. 
21: 187, 1886; Trimen & Hook. f., Handb. Fl. Ceylon 5: 33, 1900. 


Jaffna District: Jaffna, Silva 110 (PDA); Murukan, Silva in 1917 (PDA). 
Vavunia District: Ilukkulama, ca. 6 km SW of Vavunia, 7. Koyama & Herat 
13603 (NY, PDA, US). Anuradhapura District: Anuradhapura, at margin of 
Nuwara Wewa Tank, T. Koyama et al. 13939 (NY, PDA, US); Anuradhapura, 
Brodin C. P.3040 ex p. (PDA): Minneriya, ca. 15 miles E of Habarana, T. Koyama 
et al. 13568 (NY, PDA, US). Mannar District: ca. 10 miles S E of Mannar, 7. 
Koyama et al. 13922 (NY, PDA, US). Polonnaruwa District: Yoda Ela, Silva in 
1905 (PDA); Mannampitiya, ca. 5 miles ESE of Polonnaruwa, 7. Koyama & 
Herat 13572 (NY, PDA, US). Kandy District: Peradeniya, Trimen C.P. 3040 in 
part (PDA). Badula District: ca. 8 miles E of Mahiyangana, 3.5 miles N. of Mile 
53 on Mahiyangana-Padiyatalam Rd., T. Koyama et al. 13980 (NY, PDA, US). 
Trincomalee District: Trincomalee, Glenie C.P. 3788 (PDA). Batticaloa District: 
Batticaloa, Trimen C.P. 3040 in part (PDA). Monaragala District: E of south end 
of Jayanthiwewa, T. Koyama et al. 13988 (NY, PDA, US). Ratnapura District: 


130 Gardens’ Bulletin, Singapore — XXX (1977) 


Fig. 2. Cyperus pangorei Rottboell. AA, habit. B, portion of spikelet showing two glumes 
and their bases forming wings on the rhachilla. C, spikelet. D, prophyll. E, glume. F & G, 
achenes. H & J, prophylls at the base of umbel ray, Scales = 1 mm. 


Ceylon Cypereae 131 


Ratnapura, Trimen C.P. 3040 in part (PDA); Ranwala, Amaratunga 670 (PDA). 
Kurunegala District: Hettipola, Amaratunga 670 (PDA); Uhuniya, Amaratunga 
182 (PDA). Ruhuna National Park: Andunoruwa Wewa, Cooray 16 Dec. 
1969 (US). 


Distribution. Pantropic, with its range extending northwards to eastern China 
and Japan. Rare in South America, In Ceylon very common in wet places at low 
altitudes, and often noted in rice fields. 


4. Cyperus papyrus L., Sp. Pl. ed. 1, 47, 1753. 


Kandy District: cultivated in the Botanic Gardens, Peradeniya, Amaratunga 
197 (PDA). Native of East Tropical Africa with a subspecies in Madagascar. 
Widely cultivated in Ceylon as an ornamental. 


5. Cyperus corymbosus Rottboell, Descr. Icon. Rar. Nov. Pl. 42, t. 7 f. 4, 1773; 
' Thwaites, Enum. Pl. Zeyl. 344, 1864; C. B. Clarke, Fl. Brit. India 6: 612, 1893; 
Trimen & Hook. f., Handb. Fl. Ceylon 5: 29, 1900; C. E. C. Fischer in Gamble, 
Fl. Madras 9: 1641, 1931; Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 80, f. 10, 1936. 


Anuradhapura District: between Ramboda and Madawachchiya, 7. Koyama 
& Herat 13602 (NY, PDA, US). Matale District: between Naula and Dambulla, 
vicinity of Mile 40, Jayasuria 56 (US, PAD). Kandy District: Peradeniya, 
Amaratunga 199 (PDA). Kegalle District: Baddewela, T. Koyama & Samarakoon 
13546 (NY, PDA, US). Badula District: 5 miles WNW of Mahiyangana, Davidse 
7595 (MO, NY, PDA). Batticaloa District: Wandeloos Bay, T. Koyama & Herat 
13579 (NY, PDA, US). Monaragala District: E of south end of Jayanthiwewa, 
T. Koyama et al. 13987 (NY, PDA, US). Galle District: Moragoda, left bank of 
Bentota Ganga, Amaratunga 2357 (PDA). Puttalam District: Irranonillu Madampe, 
Amaratunga 2574 (PDA). 


Distribution. Tropical West Africa, Madagascar. Nepal, India, Burma, 
northern Australia, West Indies and tropical South America (Venezuela and 
Brazil). Marshy places at low altitudes. 


In Ceylon this species called Gal-éhi is occasionally more or less cultivated, 
because its straight culms are preferred by natives as material for making a kind 
of rough mats. The culms of Scirpus grossus L.f. are also used for the same purpose 
but is not cultivated. 


6. Cyperus articulatus L., Sp. Pl. ed. 1, 44, 1753; Thwaites, Enum. Pl. Zeyl. 343, 
1864; C. B. Clarke, Fl. Brit. India 6: 611, 1893; Trimen & Hook. f., Handb. FI. 
Ceylon 5: 29, 1900; C. E. C. Fischer in Gamble, Fl. Madras 9: 1641; Kiikenthal, 
Pflanzenr. 4 (20), 101 Heft: 77, 1936. 

Polonnaruwa District: Kentalai, Trimen in 1885 (PDA). Batticaloa District: 
Batticaloa, Thwaites C.P. 3561 (K, PDA). 


Distribution. Mediterranean Region, Tropical Africa, India, Indo-China, 
southeastern U.S.A. to Central and South America. More frequent in the Neotropics. 


It is rather interesting that this species has never been collected since Trimen’s 
time. 


7. Cyperus pangorei Rottboell, Descr. Icon. Rar. Nov. Pl. 31, ¢. 7 f. 3, 1773; 
Thawaites, Enum. Pl. Zeyl. 344, 1864; Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 76, 
1936. Fig. 2. 

Cyperus tegetum Roxburgh, Fl. Indica ed. 1, 1: 208, 1820; C. B. Clarke, FI. 
Brit. India 6: 613, 1893, & Illustr. ¢. 17, f. J, 1909. Cyperus dehiscens 
(Nees) Nees, Linnaea 9: 286, 1835; Kunth, Enum. Pl. 2: 56, 1837; Trimen & Hook. 
f. Handb. Fl. Ceylon 5: 30, 1900. Papyrus pangorei (Rottb.) Nees in Wight, 
Contrib. Bot. India 88, 1834, in major part, concerning basionym. Papyrus 
dehiscens Nees in Wight, l.c. 89, 1834. 


132 Gardens’ Bulletin, Singapore — XXX (1977) 


Anuradhapura District: ca. 25 miles W of Trincomalee, along Rd. A 12, 
T. Koyama et al. 14056 (NY, PDA, US); 2 km N of Yakalla, T. Koyama 13595 
(NY). Monaragala District: road between Maha Oya and Mullajama at Mile 14, 
T. Koyama et al. 13981 (NY, PDA, US); Kotagoda, Siyambaladuwa to Inginiya- 
gala, Clayton 5825 (K, PDA). Colombo District: Ja-al on Gampaha Rd., Comanor 
1026 (US). 


Distribution. Ceylon, India, Nepal and Burma. According to Kiikenthal (op. 
cit.) also cultivated in Mauritius for mat-making. 


Like C. malaccensis and C. corymbosus this species provides material for 
making mats. Trimen (op. cit.) mentions that in the Dambara area along Mahaweli 
Ganga this species is exclusively used for this purpose, and the mats are named 
after the locality. 


8. Cyperus bulbosus Vahl, Enum. Pl. 2: 342, 1806; C. B. Clarke, Fl. Brit. India 
6: 611, 1893; Trimen & Hook. f., Handb. Fl. Ceylon 5: 22, 1900; C. E. C. Fischer 
in Gamble, Fl. Madras 9: 1641, 1931; Kukenthal, Pflanzenr. 4 (20), 101 Heft: 125, 
f. [5 C-E, 1936; Kern, FI. Males. 1, 7 (3): 605, 1974. 


Puttalam District: Karapitiya, Kundu & Balakrishnan 363 (US); Chilaw, 
Ferguson, Nov. 1881 (PDA). Trincomalee District: beach S of Elizabeth Point, 
sea level, Davidse 7523 (MO, NY, PDA, US). Hambantota District: Ruhuna 
National Park, Patangala, beach, Cooray 16 Nov. 1969 (NY); Block III, 2 miles E. 
of Vaddangewardiya, N of Kataragama, Wirawan 658 (NY). 


Distribution. Tropical Africa, southern Asia, Malesia and northern Australia. 
Sandy beach; in Ceylon rather scarce. 


In southern Asia the bulbs of this species is sometimes eaten by natives. 


9. Cyperus stoloniferus Retzius, Obs. Bot. 4: 10, 1786; C. B. Clarke, Fl. Brit. 
India 6: 615, 1893 & Illustr. Cyper. t. 79 f. 1-3, 1909; Trimen & Hook. f., Handb. 
Fl. Ceylon 5: 36, 1900; Kikenthal, Pflanzenr. 4 (20), 101 Heft: 106, 1935; Blake, 
Univ. Queens]. Papers 2 (2): 9, ¢. 7, 1942; Kern, Fl. Males. I, 7 (3): 606, f. 50, 
1974. 


Jaffna District: Punkudutivu, lagoon edge, Kundu & Balakrishnan 670 (US). 
Mannar District: End of the Causeway at Mile 145, T. Koyama et al. 13923 (NY, 
PDA, US). Puttalam District: Puttalam Lagoon, ca. 6 miles N of Mampuri, T. 
Koyama et al, 13912 (NY, PDA, US); Pallugaturai. W of Wilpattu National Park, 
Davidse et al, 8217 (NY, PDA, US). Trincomalee District: Irrakkakandi, N of 
lagoon, ca. 12 miles NNW of Trincomalee, T. Koyama et al. 14067 (NY, PDA, 
US). Colombo District: Negombo, Simpson 7926 (PDA); Mount Lavinia, Trimen 
in 1981 (PDA); Kalagedihena, Amaratunga 1406 (PDA). 


Distribution. Madagascar, Mauritius, India, Indo-China, Malesia and Australia. 


In Ceylon rather frequent at margins of brackish lagoons, often forming a large 
community. 


10. Cyperus rotundus L., Sp. Pl. ed. 1, 45, 1753; Thwaites, Enum. Pl. Zeyl. 343, 
1864; C. B. Clarke, Fl. Brit. India 6: 614, 1893; Kiikenthal, Pflanzenr. 4 (20), 101 
Heft: 107, f. 13, 1935; S. T. Blake, Univ. Queensl. Papers 2 (2): 8, 4. 4, 1942; 
Kern, Fl. Males. I, 7 (3): 604, f. 49, 1974. 


; Jaffna District: Jaffna, Gardner C.P. 804, ex p. (PDA). Anuradhapura 
District: Minneriya Tank, Thwaites C.P. 804 ex p. (K, PDA); Anuradhapura, 
Sinniah in 1926 (PDA). Polonnaruwa District: Kanniyai, Ramanathan 421 
(PDA); Mile 79 on Rd. A 15, 1 m alt., Davidse 7568 (MO, NY). Kurunegala 
District: Melsiripura, Amaratunga 1333 (PDA). Matale District: Nalande, Alston 
640 (PDA). Kandy District: Peradeniya, Thwaites C.P. 804 in part (PDA); 
Kadugannawa, Alston 1089 (PDA); between Balana and Alagalla, Comanor 1187 


Ceylon Cypereae 133 


(PDA, US); Haragama, Alston in 1926 (PDA); Gannoruwa, Alston 300 (PDA); 
Katugastota, Amaratunga 1088 (PDA). 


Distribution. A cosmopolitan weed seen in tropical, subtropical and temper- 
ate regions of all the world. 


11. Cyperus tuberosus Rottboell, Descr. Icon. Rar. Nov. Pl. 28, t. 7 f. 1, 1773; 
C. B. Clarke, Fl. Brit. India 6: 616, 1893, in part. 


Cyperus rotundus L. subsp. tuberosus (Rottb.) Kiikenthal, Pflanzenr. 4 (20), 
101 Heft: 113, 1936. 


Trincomalee District: Trincomalee, Ramanathan s.n. (PDA). Polonnaruwa 
District: Polonnaruwa, Alston 292 (PDA). Mannar District: opposite to Mannar, 
Mile 145 on A 14, end of the causeway, JT. Koyama et al, 13924 (NY, PDA, US); 
along Rd. A 14, Mile 123, near the junction with Madhu Rd., 7. Koyama et al. 
13918 (NY, PDA, US). Vavuniya District: Palayanalankulan, Mile 120 on Rd. 
A 14, T. Koyama et al. 13936 (NY, PDA, US). Matale District: between Dambula 
and Habarane, at Mile 96 on Rd. A 6, T. Koyama & Herat 13563 (NY, PDA, US); 
vicinity of Mile 40, between Naula and Dambula, Jayasuria 59 (PDA). 


Distribution. Eurasia, 


12. Cyperus retzii Nees in Wight, Contrib. Bot. India 82, 1834. 


Cyperus rotundus L. subsp. retzii (Nees) Kiikenthal, Pflanzenr. 4 (20), 101 
Heft: 114, 1935. 


Central Province: without definite locality, Thwaites C.P. 3750 (PDA). 


Distribution. So far known from India, Ceylon and Australia; rare. 


Thwaites’ C.P. 3750 cited above is the only Ceylonese collection of this species. 
In this specimen relatively broad spikelets ranging from 2.8 to 3.25 mm in width 
bear lance-ovate glumes which have two closely situated parallel veins on both 
sides of the mid-nerve and attain as much as 4 mm in length. Its achenes, average 
being 1.25 mm long, tend to be slightly larger than those of C. rotundus. Because 
of these characters the identity of C.P. 3750 seems certain though I have not seen 
as yet the type of C. retzii. 


13. Cyperus scariosus R. Brown, Prodr. 216, 1810; C. B. Clarke, Fl. Brit. India 6: 
612, 1893 & Illustr. Cyper. t. 75 f. 2-3, 1909; S. T. Blake, Univ. Queensl. Papers 
2 (2): 9, t. 2, 1942: Kern, Reinwardtia 2: 103, f. 3, 1952 & Fl. Males. I, 7 (3): 
607. 1974. j 


Cyperus corymbosus Rottb. var. scariosus (R. Br.) Kiikenthal, Pflanzenr. 4 
(20), 101 Heft: 83, 1935. Cyperus mitis Steudel, Synops. Pl. Glumac. 2: 316, 
1855; Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 103, 1935; Abeywickrama, Ceylon 
Journ. Sci., Biol. Sci., 2 (2): 134, 1959. 


Jaffna District: between Poonaryn and Elephant Pass, along Jaffna Lagoon, 
T. Koyama et al. 14048 (NY, PDA, US); Jaffna, Silva in 1920 (PDA); near 
Ampan, Clayton 5235 (K, PDA); Keerimalai, Point Pedro, Clayton 5203 (K, PDA); 
Puttalam District: between Puttalam and Kurunegala, Trimen Aug. 1883 (PDA); 
Madampe, Amaratunga 2577 (PDA). Matale District: between Dambula and 
Habarane at Mile 96 on Rd. A 6, T. Koyama & Herat 13564 (NY, PDA, US); 
Dambula, Thwaites C.P. 3966 (K, PDA). Hambantota District: Tissamaharama, 
Trimen Dec. 1882 (PDA). 

Distribution. Madagascar, India to Indo-China, Malesia, Australia. 

Among very confusing nut-grasses C. scariosus can be easily recognizable by 
its filiform, obtuse-tipped spikelets which are only 1.5 to 2 mm wide. In Ceylon 
I saw this species growing together with C. tuberosus and C. rotundus respectively 
in two habitats, where I have seen no evidence of hybrids with the latter species. 


134 Gardens’ Bulletin, Singapore — XXX (1977} 


Fig. 3. Cyperus tenuiculmis Steudel. A, habit. B, spikelet. C, prophyll at the base 
of ame D, glume. E, lateral view of glume showing its hyaline extension of the base. 
F, achene 

Cyperus zollingeri steudel. G, cluster of spikelets. H, spikelet. J, portion of spikelet 
showing glumes and achenes. K, prophyll. L, glume, M, achene. = | mm. 


Ceylon Cypereae 135 


14. Cyperus sphacelatus Rottboell, Descr. Pl. Rar. Progr. 21, 1772 & Descr. 
Icon. Rar. Nov. Pl. 26, 1773. Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 129, 1935; 
Kern, Reinwardtia 2: 107, f. 5, 1952, & Fl. Males. I, 7 (3): 609, 1974. 


Kandy District: Peradeniya, Univ. of Ceylon Campus, 500 m alt., Comanor 
321 (NY, US); ca. 5 miles SE of Gampaha, Mile 18/6 on Rd. A 5, 720 m alt., 
Davidse et al. 7922 (MO, NY, PDA, US). Colombo District: Horana, Alston 
976 (PDA). 


Distribution. Tropical America and tropical Africa; introduced to Ceylon, 
Malesia, Queensland and Tahiti. 


The introduction of this species into Ceylon seems pretty old as it was already 
collected by Alston in the beginning of 1900. 


15. Cyperus tenuiculmis Bockeler, Linnaea 36: 286, 1870; Kern, Reinwardtia 3: 
30, 1954, & Fl. Males. I, 7 (3): 608, 1974. Fig. 3 A-F 


“Cyperus zollingeri Steudel”’ sensu C. B. Clarke, Fl. Brit. India 6: 613, 1893, 
& Illustr. Cyper. t. 8 f. 1-2, 1909; Trimen & Hook. f., Handb. Fl. Ceylon 5: 35, 
1900; Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 133, 1935. 

Kandy District: Hantana, Mueller-Dombois et al. June 18, 1968 (PDA). 
Nuwara-Eliya District: Ramboda, 1180 m alt., T. Koyama 13620 (NY); Hakgala, 
Mueller-Dombois et al. Jan. 1968 (PDA, US). Badulla District: ca. 8 miles E of 
Mahiyangana, 3.5 miles N of Mile 53 on Mahiyangana-Pediyatalawe Rd., T. 
Koyama et al. 13977 (NY, PDA, US); Diyatalawa, Mueller-Dombois et al. Jan. 
14, 1968 (PDA). Monaragala District: E of the south end of Jayanthiwewa Tank, 
T. Koyama et al. 13993 (NY, PDA, US). 


Distribution. Throughout the tropics of the Old World with the range extending 
northwards to southern Japan in eastern Asia. 


16. Cyperus zollingeri Steudel [in Zollinger, Syst. Verz 1: 62, 1854, name only, &] 
Synops. Pl. Glumac. 2: 17, 1855: Kern, Reinwardtia 3: 28, 1954 & Fl. Males. I, 7 
(3): 608, 1974. Fig. 3 G-M 


Cyperus ramosii Kiikenthal in Fedde, Repert. Sp. Nov. 21: 326, 1925, & 
Pflanzenr. 4 (20), 101 Heft: 136, 1935; Kern, Reinwardtia 2: 109, f. 7, 1952. 


Hambantota District, Ruhuna National Park: Yala Dunes, Ecology Project 
Plot R26, Cooray 8 Dec. 1969 (NY, PDA, US); Block I, at Rugamtota, Mueller- 
Dombois et al. 7 Mar. 1969 (NY, US); Kumbukkan Oya, ca. 2 miles above mouth, 
at Magabakanda Meda Duwa, Block II, Fosberg et al 51107 (NY, PDA, US). 


Distribution. Tropical Africa, Madagascar, Malesia, northern Australia. In 
Ceylon rare in open grassy places on sandy soil. 


New to the flora of Ceylon. This species differs from the better known 
C. tenuiculmis by the softer leaves and bracts, annual tufted habit without stolons, 
and smaller floral parts as illustrated in Fig. 3. 


17. Cyperus distans L. f., Suppl. Sp. Pl. 103, 1781; Thwaites, Enum. Pl. Zeyl. 344, 
1864; C. B. Clarke, Fl. Brit. India 6: 607, 1893; Trimen & Hook. f., Handb. FI. 
Ceylon 5: 30, 1900; Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 137, 1935; Kern, FI. 
Males. I, 7 (3): 610, 1974. 


Polonnaruwa District: W of Siva Devale in Sacred Area Section 2A, Ripley 
369 (PDA). Kegalle District: Kadugannawa, Grupe 2/3 (PDA, US). Nuwara 
Eliya District: Hakgala Botanic Garden, Clayton 5784 (K, PDA); Maturata, 
Ferguson in 1906 (PDA). Kandy District: Peradeniya, Thwaites C.P. 810 ex p. 
(K, PDA); Gannoruwa, Alston 319 (PDA); ca. 2 miles E. of Maddakele, slope 
N.W. of Kunckles Peak, 1440 m alt., Davidse 8320 (MO, NY, PDA, US). 
Batticaloa District: Batticaloa, Thwaites C.P. 810 in part (PDA). 


Distribution. Pantropic. 


136 Gardens’ Bulletin, Singapore — XXX (1977) 


18. Cyperus nutans Vahl, Enum. PI. 2: 363, 1806; C. B Clarke, Fl. Brit. India 6: 
607, 1893; Trimen & Hook. f., Handb. Fl. Ceylon 5: 31, 1900; C. E. C. Fischer in 
Gamble, Fl. Madras 9: 1640; 1931; Kukenthal, Pflanzenr. 4 (20), 101 Heft: 144, 
f. 5 A-D, 1935; Kern, Fl. Males. I, 7 (3): 609, 1974. 


Cyperus distans L. var. major Thwaites, Enum, Pl. Zeyl. 432, 1864. 


18a. Subsp. nutans. 


Anuradhapura District: Anuradhapura, Thwaites in 1881 (PDA); Wilpattu 
National Park, ca. 2.7 miles to Ranger’s Office along pass from southern entrance, 
T. Koyama & Jayasuria 13967 (NY, PDA, US). Trincomalee District: Trinco- 
malee, Brodie C.P. 3844 (PDA). Matale District: Dambulla, Thwaites C.P. 3966 in 
part (PDA). Colombo District: Galapitamada, Amaratunga 1148 (PDA). Monara- 
gala District: E of south end of Jayanthiwewa Tank, T. Koyama et al. 13992 (NY, 
PDA, US). Hambantota District: Ruhuna National Park, Yala Plain, Comanor 
863A (NY, US). 


Distribution. India. southern China and Malesia. 


18b. Subsp. eleusinoides (Kunth) T. Koyama, stat. nov. 


Cyperus eleusinoides Kunth, Enum. Pl, 2: 29, 1937; C. B. Clarke, FI. Brit. 
India 6: 608, 1893; Kiukenthal, Pflanzenr. 4 (20), 101 Heft: 144, 1936. 
Cyperus nutans Vahl var. eleusinoides (Kunth) Haines, Bot. Bihar Orissa 5: 898, 
1924: Kern, Fl. Malesiana I, 7 (3): 610, 1974. 


Kandy District: Haragama, Trimen C.P. 3044 (PDA). Hambantota District: 
Ruhuna National Park, Mueller-Dombois et al. Jul. 1967 (PDA). 


Distribution. Tropical Africa, India to Malesia and northern Australia, also 
extending northwards to the Ryukyus. 


Cyperus nutans and C. eleusinoides are very similar to one another, and the 
only reliable difference between the two is the more densely disposed, longer 
spikelets in the latter. But, this difference becomes obscure as ssp. subprolixus of 
the Ryukyus and Taiwan comes between the two as to the density and the number 
of florets in the spikelet. I, therefore, regard C. eleusinoides as a subspecies of 
C. nutans. 


19. Cyperus procerus Rottboell, Descr. Icon. Rar. Nov. Pl. 29, ¢. 5 f. 3, 1773; 
Thwaites, Enum. PJ. Zeyl. 343, 1864; C. B. Clarke, Fl. Brit. India 6: 610, 1893; 
Trimen & Hook. f., Handb. Fl. Ceylon 5: 34, 1900; Kiikenthal, Pflanzenr. 4 (20), 
101 Heft: 91, 1935; Kern, Fl. Males. I, 7 (3): 611, f. 51, 1974. Fig. 4 


Jaffna District: Murunkan, Silva in 1917 (PDA). Vavunia District: Vavunia, 
Clayton 5306 (K, PDA); near Kokkavil, Clayton 5292 (K, PDA). Anuradhapura 
District: Anuradhapura, Trimen Dec. 1884 (PDA); Alankulam, Trimen Oct. 1883 
(PDA); Mile 123 on Rd. A 14, E of Medachchiya, T. Koyama et al. 13921 (NY, 
PDA, US). Wilpattu National Park: Kuda Pathessa, T. Koyama & Herat 13394 
(NY, PDA, US); Periya Naga Villu, Wirawan et al. 903 (US); Manikepola Uttu, 
T. Koyama 13461 (NY, PDA, US); Wilpattu West Sanctuary, Mari Villu, 7. 
Koyama 13972 (NY, PDA, US). Batticaloa District: Batticaloa, Thwaites C.P. 
3752 (K, PDA); Ottawady, opposite to Valaichchenai, T. Koyama & Herat 13582 


(NY, PDA, US). Mannar District: Murunkan, Silva in 1917 (PDA); Illupadichcha- 


nai, Lord s.n. (PDA). Puttalam District: Madampe, Amaratunga 2575 (PDA). 
Matale District: Dambulla, Thwaites C.P. 3752 in part (K, PDA); between Naula 
and Dambula, vicinity of Mile 40, Jayasuria 60 (PDA). Kandy District: Siyam- 
balatenna, Alston 390 (PDA); Peradeniya, Thwaites C.P. 802 (PDA), Alston 
118I (PDA). Kurunegala District: Kurunegala, Thwaites C.P. 3752 in part (PDA). 
Colombo District: Colombo, Ferguson s.n. (PDA). Hambantota District: Tissa- 
maharama, Alston 1179 (PDA); Ruhuna National Park, Block II, 7 m alt. 


Ceylon Cypereae 137 


\ 
Sra 44 
RK 

=e 


we 
SS 


Fig. 4. Cyperus procerus Rottboell. AA, habit. B, spikelet. C, prophyll. D, glume. 
E, achene. Scales for floral parts = | mm. 


138 Gardens’ Bulletin, Singapore — XXX (1977) 


Comanor 1037 (PDA), US). Amparai District: Helawe Eliya, ca. 7 miles S. of 
Panama, E. of Helawe Lagoon, 7. Koyama et al. 14024 (NY, PDA, US). 


Distribution, India, Ceylon, Indo-China, Taiwan, Malesia, Australia (Queens- 
land). In Ceylon abundant in marshy places at low altitude. 


20. Cyperus pilosus Vahl, Enum. Pl. 2: 354, 1806; Thwaites, Enum. Pl. Zeyl. 344, 
1864; C. B. Clarke, Fl. Brit. India 6: 609, 1893; Trimen & Hook. f., Handb. FI. 
Ceylon 5: 32, 1900; Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 92, 1935; Kern, FI. 
Malesi I, 7 (3): 611, f. 52, 1974. 


Cyperus marginellus Nees in Wight, Contrib. Bot. India 83, 1834. 
Cyperus obliquus Nees in Wight, I.c. 86, 1834. 


Nuwara Eliya District: between Pussellawa and Ramboda, Mile 32/8 on 
Nuwara-Eliya-Kandy Rd., 1000 m alt., 7. Koyama 13617 (NY, PDA, US). Badulla 
District: Haptale, Silva in 1906 (PDA). Kandy District: Rangala, Ferguson Sept. 
1885 (PDA). Kalutara District: Korosduwa, Amaratunga 2541 (PDA); Kande 
Kanda, Amaratunga 2523 (PDA). Kegalle, District: Kegalle, Amaratunga 1651 
(PDA); Kattiyakumbura, Amaratunga 1589 (PDA); Nayinankada, 700 m alt., 
T. Koyama & Samarakoon 13561 (NY, PDA, US). Kurunegala District: Nara- 
munala, Amaratunga in 1966 (PDA); Malsiripura, Amaratunga 1330 (PDA). 
Ratnapura District: 11 miles E of Deniyaya at Mile 62 on Rd. A 17, 750 m alt., 
Davidse 7876 (MO, NY, PDA, US). Galle District: Akeemana, Alston 1187 
(PDA); Moragoda, Amaratunga 2359 (PDA). Monaragala District: Inginiyagala 
National Park, between Baduluwila and ““Westminster Abbey”, T. Koyama et al. 
13994 (NY, PDA, US); E of south end of Jayanthiwewa, T. Koyama et al. 13988 
(NY, PDA, US). 


Distribution. Rather widely distributed in tropical and subtropical regions of 
the Old World from tropical West Africa through Central Asia and Indian Sub- 
Continent eastwards to Malesia and northern Australia, In eastern Asia the range 
extending northwards as far as to warm-temperate Japan. 


Although this species is highly variable in the dimension of leaves and inflo- 
rescences, it is well demarcated by the subdensely hispid rhachis of spikes. 


21. Cyperus compressus L., Sp. Pl. ed. 1, 46. 1753; Thwaites, Enum. Pl. Zeyl. 342, 
1864; C. B. Clarke, Fl. Brit. India 6: 605, 1893; Trimen & Hook. f., Handb. FI. 
Ceylon 5: 33. 1900; Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 156, f. 4 A-D, 1935; 
Kern, Fl. Males. I, 7 (3): 617,1974. 


21a. Subsp. compressus. 


Jaffna District: Jaffna, Trimen Feb. 1890 (PDA), Silva in 1920 (PDA). 
Vavunia District: S of Mullaittivu, Fosberg & Balakrishnan 53527 (NY, US). 
Polonnaruwa District: Gal Oya Reservoir, near the spilway, 270 m alt., Comanor 
561b (US). Trincomalee District: 7 miles S of Dambulla on Kandy Rd., Clayton 
5078 (K, PDA); between Naula and Dambulla, vicinity of Mile 40, Jayasuria 57 
(PDA). Kurunegala District: Kurunegala, Thwaites C.P. 812 (K, PDA). Kandy 
District: Peradeniya, Trimen C.P. 812 (PDA); Kandy, Alston in 1926 (PDA). 
Colombo District: Danowita, Amaratunga 1450 (PDA). Hambantota District: 
Ruhuna National Park, Gonalabbe, Block I, Cooray & Balakrishnan 20 Jan. 1969 — 
(US). Monaragala District: E of the south end of Jayanthiwewa, T. Koyama et al. 
13991 (NY, PDA, US); way to Nilgola, Trimen Jan. 1888 (PDA); Inginiyagala, 
Amaratunga 1706 (PDA). 


Distribution, Cosmopolitan; tropical, subtropical and temperate regions of all 
world; weed of cultivated and waste places, more frequent along sea coast. 


Ceylon Cypereae 139 


21b. Subsp. micranthus [. Koyama, subsp. nov. 


Planta omnibus partibus minor; inflorescentia semper in capitulum unicum 
congesta; glumae 2 mm longae; achaenia obovato-orbicularia, basi valde attenuantia, 
vix 1 mm longa, 0.75 mm lata; caeteroquin sicut ssp. compressus. 


Trincomalee District: exposed eastern bottom of Kantalai Tank at Mile 134 
on Habarana-Trincomalee Rd., Davidse 7561 (NY, holotype; MO, isotype). 


Distribution. Thus far known only by the collection cited above. 


Subspecies micranthus differs from the typical phase primarily by the smaller 
glumes (2 mm vs. ca. 3.5 mm in length) and achenes (1 by 0.75 mm ys. 1.5 by 
1 mm in size) in addition to its comparatively smaller habit with always congested 
inflorescences. Besides the fertile achenes every detail of floral parts shows no 
evidence of the possible hybrid nature of this subspecies, though it has been known 
that C. compressus rather freely hybridize with several species. So far I have not 
come across as yet any specimen of C. compressus bearing such smaller glumes 
and achenes from its total range. 


22. Cyperus iria L., Sp. Pl. ed. 1, 45, 1753; Thwaites, Enum. Pl. Zeyl. 344, 1864; 
C. B. Clarke, Fl. Brit. India 6: 606, 1893; Trimen & Hook. f., Handb. Fl. Ceylon 
5: 18, 1900; Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 150, 1935; Kern, Fl. Males. I, 
7 (3): 616, 1974. 


Jaffna District: Jaffna, Silva Nov. 1920 (PDA). Trincomalee District: Trinco- 
malee, Ramanathan Jul. 1926 (PDA). Anuradhapura District: Mile 123 on Rd. 
A 14, T. Koyama et al. 13920 (NY, PDA, US); Wilpattu National Park, Manike- 
pola Uttu, T. Koyama & Jayasuria 13951 (NY, PDA, US). Matale District: between 
Mirisgoniyawa and Dambulla, Silva Nov. 1926 (PDA); between Naula and Dam- 
bulla, vicinity of Mile 40, Jayasuria 55 (PDA); Matale North, Jayasuria 53 (PDA). 
Kandy District: Peradeniya, Trimen C.P. 811 (K, PDA); Pundaluoya, Hughes 
Dec. 1902 (PDA). Kurunegala District: Kurunegala, Thwaites C.P. 811 in part 
(PDA). Colombo District: Giriulla, Amaratunga 998 (PDA); Veyangoga, Amara- 
tunga 1946 (PDA). Kalutara District: Bolgoda, Amaratunga 2524 (PDA). Ruhuna 
National Park: Buttawa Plain, near Ecology Project Plot R35, Cooray 12 Dec. 
1969 (US); Yala Plain, Patanagala Beach, 3 m alt., Comanor 863B (NY, US). 


Distribution. Widely distributed in Asia from Iran and Afghanistan through 
Indian Subcontinent northeastwards to China and Japan, and eastwards to Malesia 
and Australia; also in East Africa. Apparently introduced to southeastern U.S.A., 
West Indies and South America. Common weed in cultivated ground and rice field. 


23. Cyperus diffusus Vahl, Enum. Pl. 2: 321, 1806; C. B. Clarke, Fl. Brit. India 
6: 603, 1893; Trimen & Hook. f., Handb. Fl. Ceylon 5: 28, 1900 Kikenthal, 
Pflanzenr. 4 (20), 101 Heft: 208, 1936; Kern, Fl. Males. I, 7 (3): 619, 1974. 


Cyperus nigro-viridis Thwaites, Enum. Pl. Zeyl. 344, 1864. 


23a. Subsp. diffusus. 
Kurunegala District: Kurunegala, Thwaites C.P. 2879 (PDA). 
Distribution. India, Indo-China, southern China and Malesia. 


23b. Subsp. macrostachyus (Boéckleler) T. Koyama, stat. nov. 


Cyperus diffusus Vahl var. macrostachyus Béckeler, Linnaea 35: 534, 1868; 
Trimen & Hook. f., Handb. Fl. Ceylon 5: 28, 1900; Kiikenthal, Pflanzenr. 4 (20), 
101 Heft: 209, 1936; Kern, Fl. Males. I, 7 (3): 619, f. 54, 1974. Cyperus 
pubisquama Steudel [in Zollinger, Syst. Verz. 1: 62, 1854, nomen,] Synops. PI. 
Glumac. 2: 20, 1855. 


140 Gardens’ Bulletin, Singapore — XXX (1977) 


Kurunegala District: Kurunegala, Thwaites C.P. 3931 (PDA); Mallawapitiya, 
Amaratunga 1101 (PDA); Athagala, Amaratunga 678 (PDA). Matale District: 
Matale, Ferguson Oct. 1884 (PDA). 


Distribution. From India through Indo-China eastwards to Malesia. 


Subspecies macrostachyus differs from the typical phase in its generally larger 
habit, far more copious inflorescence and longer spikelets as mentioned in the key 
to species. In Ceylon subsp. diffusus is very rare, and has so far been documented 
only by Thawaites C.P. 2879. 


24. Cyperus platystylis R. Brown, Prodr. Fl. Nov. Holl. 214, 1810; C. B. Clarke, 
Fl. Brit. India 6: 598, 1893; Trimen & Hook, f., Handb. Fl. Ceylon 5: 24, 1900; 
C. E. C. Fischer in Gamble, Fl. Madras 9: 1639, 1931; Kiikenthal, Pflanzenr. 
4 (20), 101 Heft: 185, f. 27, 1936; Kern, Fl. Males. I, 7 (3): 618, 1974. 
Fig. 5 

Cyperus pallidus Nees [Linnaea 9: 284, 1835, nomen nudum, &] in Wight, 
Contrib. Bot, India 79, 1834; Thwaites, Enum. Pl. Zeyl. 34, 1864. Not of 
Willdenow ex Link, 1820, nor of Savi, 1830. Anosporum pallidum (Nees) 
Bockeler, Linnaea 36: 412, 1870. 


Kurunegala District: Kurunegala, Trimen C.P. 3559 in part (PDA); Wariya- 
pola, Amaratunga 1104 (PDA). Batticaloa District: Kalmunai, Ahmed Aug. 1940 
(PDA); Batticaloa Trimen C.P. 3559 in part (K, PDA). Amparai District: Lahu- 
gala Tank, T. Koyama et al. 14030 (NY, PDA, US). Colombo District: Mutu- 
rajawela, Amaratunga 144 (PDA). 


Distribution. India and Ceylon to Taiwan, and through Malesia eastwards to 
northern and eastern Australia. In marshes and ponds usually growing in large 
communities, 


This rare species is easily discernible in its achenes with spongy thickened 
angles and densely spiculose hemisphaerical umbel. The rhizomes are often more 
or less elongated and obliquely ascending, but do not emit stolons. 


25. Cyperus alternifolius L., Mant. 2: 28, 1771. 

This species, known from East Africa and Madagascar, is represented in 
Ceylon by the following. 
Subsp. flabelliformis (Rottboell) Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 193, 1936. 


Cyperus flabelliformis Rottboell, Descr. Pl. Rar. Progr. 22, 1772, & Descr. 
Icon. Rar. Nov, Pl..-42, f° 92°." 2) 14.773. 


Badulla District: Rawanaella Waterfall, ca. 2 miles SE of Ella, Mile 13, 
780 m alt., Davidse et al. 8858 (MO, NY, PDA, US). Monaragala District: 4 
miles N of Wellawaya, Wheeler 12689 (PDA, US). 


Distribution. A native of tropical Africa and Arabia. Widely cultivated as an 
ornamental; in Ceylon escaped and established in grassy places. 


26. Cyperus radians Nees & Meyen [ex Nees, Linnaea 9: 285, 1835, nomen 
nudum,] ex Kunth, Enum. Pl. 2: 95, 1837, as “‘radicans”; C. B, Clarke, Fl. Brit. 
India 6: 605, 1893; Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 214, 1936; Kern, FI. 
Males. I, 7 (30): 623, 1974. 


Trincomalee District: seashore several miles E of Toppur, ca. 15 miles SE of 
Trincomalee, 7. Koyama 14060b. (NY). 

Distribution. Indo-China, Malay Peninsula, southeastern China, Borneo. 
Sporadically noted in sandy sea coast. 


New to the flora of Ceylon. The above collection marks a considerable range 
extension of this southeastern Asian species into the Indian Subcontinent. This is a 


Ceylon Cypereae 141 


SSS Es ©, je 


44 Se es KA AT 
\ 2s es —_ <a 


-— 

ad i 
ive d 
m3 


~ ~ a 
ss 


ZF 


SIRENS 
LAN G Q 


] 


Fig. 5. Cyperus platystylis R. Brown. AA, habit. B, spikelet. C, glume. D, dorsal view 
of achene. E, ventral side of achene. Scales = | mm, 


142 Gardens’ Bulletin, Singapore — XXX (1977) 


peculiar looking species growing in sand. Since the culms are normally extremely 
abbreviated and hidden in the leaf tuft, the elongated umbel rays look like tufted 
culms. 


27. Cyperus pulcherrimus Willdenow ex Kunth, Enum. Pl. 2: 1837; C. B. Clarke, 
Fl. Brit. India 6: 600, 1893, & Illustr. Cyper. t. 10 f. 1-3, 1909; Trimen & Hook. 
f., Handb. Fl. Ceylon 5: 27, 1900; Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 242, 
1936; Kern, Fl. Males. I, 7 (3): 624, 1974. 


“Cyperus silletensis Nees’ sensu Thwaites, Enum. Pl. Zeyl. 343, 1864. 


Wilpattu National Park: between Kattankandal Kulam and Occapu Junction, 
Wirawan et al. 977 (US); Manikepola Uttu, T. Koyama et al. 13462 (NY, PDA, 
US); between Kunbuk Wila and Kokkare Villu, T. Koyama & Jayasuria 13946b 
(NY); Malimaduwa Guards Quater, Wirawan & Cooray 1141 (NY, PDA). 
Batticaloa District: Batticaloa, Trimen C.P. 3558 (K, PDA). Kurunegala District: 
Kurunegala, Rasanayaka in 1927 (PDA). Matale District: 8 miles E, of Habarane, 
T. Koyama & Herat 13567 (NY, PDA, US). Colombo District: Kotugoda, 
Amaratunga 2027 & 2262 (PDA). 


Distribution. India, Indo-China, Malesia. 


28. Cyperus haspan L., Sp. Pl. ed. 1, 45, 1753; Thwaites, Enum. Pl. Zeyl. 343, 
1864; C. B. Clarke, Fl. Brit. India 6: 600, 1893; Trimen & Hook. f., Handb. FI. 
Ceylon 5: 26, 1900; Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 247, f. 28 E-G, 1936; 
Kern, Fl. Males. I, 7 (3): 624, ff. 56 & 57, 1974. 


Anuradhapura District: Anuradhapura, Brodie C.P. 799 (K, PDA); Wilpattu 
National Park, Manikepola Uttu, T. Koyama & Jayasuria 13953 (NY, PDA, US). 
Trincomalee District: Trincomalee, Glenie C.P. 799 in part (PDA); Periya Kulam, 
ca. 7 miles NW of Trincomalee, 7, Koyama et al. 14066 (NY, PDA, US). Puttalam 
District: Wilpattu West Sanctuary, Marai Villu, 7. Koyama 13973 (NY, PDA, 
US); Madape, Amaratunga 2569 (PDA). Vavunia District: Kokkavil, Clayton 
5294 (K, PDA). Nuwara Eliya District: Rangalla to Corbett’s Gap, Ballard 1416 
(K, NY). Kurunegala District: Kurunegala, Rasanayake in 1927 (PDA); Ibbaga- 
muwa, Amaratunga 1581 (PDA). Kandy District: Peradeniya, Alston 311 (PDA); 
Upper Hantana Rd., Comanor 314 (PDA, US); Hantane, Thwaites C.P. 965 in 
part (K, PDA). Kegalle District: Alapalawala, along river Watura Oya, ca. 500 m 
alt.. T. Koyama & Samarakoon 13558 (NY, PDA, US); Mawanella, Amaratunga 
1199 (PDA). Badulla District: Rd. A 4, ca. 3 miles W of Koslanda, ca. 900 m alt., 
T. Kvyama et al, 14032 (NY, PDA, US). Ratnapura District: 11 miles E of 
Deniyana at Mile 62 on Rd. A 17, 750 m alt., Davidse 7885 (MO, NY, PDA, US); 
Kuruwita, Trimen in 1895 (PDA). Colombo District: Muthurajawela, Amaratunga 
135 (PDA). Kalutara District: Nugegoda, Bolgoda Lake Scheme, Amaratunga 
2538 (PDA). Galle District: Galle, Thwaites C.P. 965 (PDA): Bentota Ganga, 
Amaratunga 2332 (PDA); Ambalangoda, Amaratunga 2642 (PDA). Monaragala 
District: ca. 3 miles W. of Wellawaya at Mile 135/10 on Rd. A 4, 990 ft. alt., 
Davidse 7731 (MO, NY). 


Distribution. Tropical and subtropical regions of all the world with its range 
extending northwards into Japan in East Asia. Abundant in wet places; frequent 
in rice fields. 


This species is sometimes divided into two subspecies, ssp. haspan and ssp. 
junciformis Kiikenthal, the latter differing from the typical phase in having long- 
creeping rhizome along which the culms are disposed in a row in a more or less 
spaced manner. In plants from Asia these two subspecies seem quite well circum- 
scribed, and consequently, appear to be recognizable. Typical ssp. haspan, with 
the culms tufted without conspicuous rhizome, is often confused with closely related 
C. tenuispica, The only reliable character to separate these two are floral glumes. 
In C. tenuispica the floral glumes are spaced exposing a considerable part of the 


Ceylon Cypereae 143 


subtending achene between two glumes, while in C. haspan the achenes are com- 
pletely hidden by the closely disposed glumes. 


29. Cyperus tenuispica Steudel, Synops. Pl. Glumac. 2: 11, 1855: Kiikenthal, 
Pilanzenr. 4 (20), 101 Heft: 245, f. 28 A-D, 1936; Kern, Fl. Males. I, 7 (3): 625, 
f. 58, 1974. 


“Cyperus flavidus Retzius” sensu C. B. Clarke, Fl. Brit. India 6: 600, 1893; 
Trimen & Hook. f., Handb. Fl. Ceylon 5: 27, 1900. 


Jaffna District: Mullaitiva, Ferguson s.n. (PDA); south shore of Jaffna Lagoon, 
ca. 8 miles SE of Poonaryn, T. Koyama et al. 14046 (NY, PDA, US). Wilpattu 
National Park; Kuda Pathessa, 7. Koyama 13387 (NY). Matale District: Matale 
north, Jayasuria 54 (PDA). Kandy District: Paradeniya, Alston in 1926 (PDA), 
Trimen C.P. 805 (PDA); Gannoruwa, Alston 315 (PDA). Colombo District: 
Gampaha, Simpson 8602 (PDA); Makawita, Amaratunga 1779 (PDA). Hamban- 
tota District: Ruhuna National Park, Block I, Rakinawala, Cooray 7 Dec. 1969 
(US). 


Distribution. Tropical Africa, India, Nepal, Indo-China, Malesia, south- 
eastern China, Japan. Wet places; often as a weed in rice field. 


30. Cyperus difformis L., Cent. Pl. 2: 6, 1756; C. B. Clarke, Fl. Brit. India 6: 599, 
1893; Trimen & Hook. f., Handb. Fl. Ceylon 5: 25, 1900; Kiikenthal Pflanzenr. 
4 (20), 101 Heft: 237, f. 27 F-H, 1936; Kern, Fl. Males. I, 7 (3): 629, 1974. 


Trincomalee District: Trincomalee, Ramanathan in 1926 (PDA). Polonaruwa 
District: ca. 1 mile E of Kolakanaweli, at Mile 56/7 on Rd. A 11, 7. Koyama 
13577 (NY). Batticaloa District: Batticaloa, Thwaites C.P. 3042 (K, PDA). Kandy 
District: Gannoruwa, Alston 316 (PDA). Matale District: Ereula Tank, ca. 5 
miles SE of Dambulla, 650 ft. alt., Davidse 7390 (MO, NY). Colombo District: 
Danowita, Amaratunga 192 (PDA). Amparai District: Tandiadi Kalapu Lagoon, 
S of Mile 215 on Rd. A 4, T. Koyama et al. 14015 (NY, PDA, US). Ruhuna 
National Park: Komawa Wewa, Cooray 10 Dec. 1969 (NY), US); Uraniya, 
Cooray 22 Mar. 1970 (NY). 


Distribution, Widely distributed in Eurasia from southern Europe through 
India and China to Japan and Malesia, also in Pacific Islands and Australia; 
introduced to Central America and South Africa possibly with rice. 


31. Cyperus castaneus Willdenow, Sp. Pl. 1: 278, 1797; Thwaites, Enum. Pl. Zeyl. 
343, 1864; C. B. Clarke, FI. Brit. India 6: 598, 1893; Trimen & Hooks. f., Handb. 
FI. Ceylon 5: 25, 1900; Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 264, 1936; Kern, FI. 
Males. I, 7 (3): 630, 1974. 


Anuradhapura District: Wilpattu National Park, East Intermediate Zone, 
along Wildlife Dept. Path, ca. 2 km from its barrier gate. T. Koyama & Jayasuria 
13966 (NY, PDA, US). Polonnaruwa District: SE of Giritale Wewa near Circuit 
Bangalow, Fosberg et al. 51955 (PDA, US); Polonnaruwa, Clayton 5115 (K, 
PDA). Matale District: Dambulla Rock, Trimen in 1896 (PDA). Kegalle District: 
Simpson 8358 (PDA). Badulla District: between Ratupahana and Haldunmulla, 
Ormiston in 1909 (PDA). Ruhuna National Park: Block I, Cooray 7 Dec, 1969 
(US); between Buttawa and Karangaswela, Cooray 3 Dec. 1969 (NY, US). 


Distribution. From India through Indo-China to Malesia and northern 
Australia. 


32. Cyperus cuspidatus Kunth in Humb., Bonpl. & Kunth, Nov. Gen. et Sp. 
Pl. 1: 204. 1815: C. B. Clarke, FI. Brit. India 6: 598, 1893; Trimen & Hook. f., 
Handb. Fl. Ceylon 5: 26, 1900; Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 261, 
f. 29 E, 1936; Kern, Fl. Males. I, 7 (3): 629, f. 59, 1974. 


Cyperus angustifolius Nees in Wight, Contrib. Bot. India 79, 1834. 


144 Gardens’ Bulletin, Singapore — XXX (1977) 


Anuradhapura District: Wilpattu National Park NE of Kuruttu Pondi Villu, 
T. Koyama 13413 (NY). Polonnaruwa District: 7 miles NW of Polonnaruwa, SE 
of Giritale Wewa, Fosberg & Ripley 51955 (US). Nuwara Eliya District: ca. 15 
miles NW of Nuwara Eliya, at Mile 32/8 on Road A 5, 1020 m alt., Davidse 
et al. 7946 (MO, NY, PDA). Monaragala District: Inginiyagala National Park, 
between Baduluwila and ‘‘Westminster Abbey,” 7. Koyama et al. 13997 (NY, 
PDA, US); ca. 25 miles W of Pottuvil on road to Wellawaya, near Mile 76, 
Davidse et al. 8934 (MO, NY, PDA). Hambantota District: Ruhuna National 
Park, Karasugaswela, Cooray 12 Dec. 1969 (NY, PDA). 


Distribution. Pantropic, with its range extending north to southern China and 
Formosa. 


33. Cyperus arenarius Retzius, Obs. Bot. 4: 9, 1786; Nees in Wight, Contrib. Bot. 
India 77, 1834; Thwaites, Enum. Pl. Zeyl. 342, 1864; C. B. Clarke, FI. Brit. India 
6: 602, 1893; Trimen & Hook. f., Handb. Fl. Ceylon 5: 23, 1900; C. E. C. Fischer 
in Gamble, Fl. Madras 9: 1640, 1931; Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 270, 
1936. Fig. 6 

Bobartia indica L., Fl. Zeyl. 17, 1747. — Cyperus conglomeratus Rottb. 
var. arenarius (Retzius) Cosson, Explor. Algir. Bot. 2: 245, 1855. 

Mannar District: Talaimannar, Silva in 1916 (PDA). Trincomalee District: 
Trincomalee, Glenie C.P. 798 (K, PDA); Puttalam District: Kalpitiya, Trimen in 
1883 (PDA); Chilaw, Trimen in 1880 (PDA); Wilpattu National Park, West 
Intermediate Zone, Pallugaturai Beach, Wheeler 12104 (PDA). Batticaloa District: 
Passikudah, Mueller-Dombois in 1968 (PDA); Batticaloa, Gardener C.P. 798 
(PDA). Colombo District: Colombo, Ferguson C.P. 794 (PDA); Uswetakeiyawa, 
Amaratunga 126 (PDA). Amparai District: Pottuvil, Rest House beach, Bala- 
krishnan 389 (PDA); Arugam Bay, T. Koyama et al. 14029 (NY, PDA, US), 
Fosberg & Sachet 53043 (NY, US). Hambantota District: Bentota, beach behind 
rest house, Ballard 1510 (K, PDA); Ruhuna National Park, beach E of Buttawa 
Modera, 2-3 m alt., Fosberg 50315 (US); Patanagala Beach, Cooray 17 Nov. 1969 
(PDA, US). 

Distribution. Southern Iran, Pakistan, India, Ceylon, Cochinchina. 


A coastal species easily reconginable by the whitish head with broad spikelets 
and extensive rhizome system, with which it usually forms a large pure community. 


34. Cyperus conglomeratus Rottboell, Descr. Icon. Rar. Nov. Pl. 21, t J5 f.7, 
1773; Thwaites, Enum. Pl. Zeyl. 343, 1864; C. B. Clarke, Fl. Brit. India 6: 602, 
1893; Trimen & Hook. f., Handb. Fl. Ceylon 5: 23, 1900; Kiikenthal, Pflanzenr. 
4 (20), 101 Hen: 27 2..3c Fos leo. 


Cyperus pachyrrhizus Nees ex Bockeler, Linnaea 35: 545, 1868; C. B. Clarke, 
Fl. Brit. India 6: 603, 1893; C. E. C. Fischer in Gamble, Fl. Madras 9: 1640, 
193 t: Cyperus conglomeratus Rottb. var. pachy[r]rhizus (Nees ex 
Bockeler) Trimen in Trimen & Hook. f, Handb. Fl. Ceylon 5: 23, 1900. 


Puttalam District, Wilpattu National Park: Kollankanatta Beach, Cooray 28 
Sept. 1969 (US); Pallugaturai, 7. Koyama & Jayasuria 13960 (NY, PDA, US), 
Fosberg et al. 50917 (NY, US), Wheeler 12105 (PDA). 

Distribution. Mediterranean Region, North Africa, East Africa, Iran, Arabia, 
Madagascar, southern India and Ceylon. 

This is one of the western Asian element reaching Ceylon, Having compared a 
good series of Indian and Ceylonese specimens with those from western Asia I am 
unable to separate Indian plants from western Asian ones. 


35. Cyperus cephalotes Vahl, Enum. PI. 2: 311, 1806; C. B. Clarke, FI. Brit. India 
6: 597, 1893, & Illustr. Cyper. t. 6 f. I-7, 1909; Trimen & Hook. f., Handb, FI. 
Ceylon 5: 17, 1900; Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 292, 1936; Kern, FI. 
Males. I, 7(3): 633, ff. 61-62, 1974. 


ee - 


145 


Ceylon Cypereae 


Fig. 6. Cyperus arenarius Retzius. A, habit. B, spikelet. C, prophyll. D, glume. E, pistil 
F, achene. Scales = 1 mm. 


146 Gardens’ Bulletin, Singapore — XXX (1977) 


Cyperus hookerianus Thwaites, Enum. Pl. Zeyl. 342, 1864. Anos- 
porum cephalotes (Vahl) Kurz, Journ. Asiat. Soc. Bengal 45 (2): 159, 1876. 


Kurunegala District: Ibbagamuwa, Amaratunga in 1967 (PDA). 


Distribution. From India through Indo-China northeastwards to southern 
China, and eastwards to Malesia and northern Australia. 


Trimen (op. cit.) reported this species from Galle, Lunugala and Uva. 
Currently in the Peradeniya Herbarium the specimen cited above is the only sheet. 


36. Cyperus alopecuroides Rottboell, Descr. Pl. Rar. Progr. 20, 1772, & Descr. 
[con. Rar. Nov. Pl. 38, ¢. 8 f. 2, 1773; Thwaites, Enum. Pl. Zeyl. 342, 1864; 
Trimen & Hook. f., Handb. Fl. Ceylon 5: 38, 1900; Kiikenthal, Pflanzenr. 4 (20), 
101 Heft: 71, 1935; Kern, Fl. Males. I, 7 (3): 603, 1974 


Juncellus alopecuroides (Rottb.) C. B. Clarke, Fl. Brit. India 6: 595, 1893. 


Anuradhapura District: between Ratmale and Talawa, ca. 7 miles SSW of 
Anuradhapura, T. Koyama & Herat 13608 (NY, PDA, US). Polonnaruwa District: 
Minnariya Tank, Trimen in 1884 (PDA). Batticaloa District: Batticaloa, Trimen 
C.P. 3560 (K, PDA). Hambantota District: Tissamaharama, Trimen in 1882 
(PDA). 


Distribution. North and Tropical Africa, Madagascar, India, Indo-China, 
Malesia and northern Australia. 


37. Cyperus pygmaeus Rottboell, Descr. Icon. Rar. Nov. Pl. 20, ¢. 14 f. 4-5 
1773; Nees in Wight, Contrib. Bot. India 72, 1834; Trimen & Hook. f., Handb. 
Fl. Ceylon 5: 18, 1900; Kern, Fl. Males. I, 7 (3): 634, 1974. 


Juncellus pygmaeus (Rottb.) C. B. Clarke, Fl. Brit. India 6: 596, 1983. 
Cyperus michelianus (L.) Delile subsp. pygmaeus (Rottb.) Aschers, & 
Graebn., Synops. Mitteleur. Fl. 2 (2): 273, 1903; Kiikenthal, Pflanzenr. 4 (20), 101 
Heft: 312, f. 35 F-G, 1936. 


Polonnaruwa District: Gal Oya Reservoir, near spilway, 270 m alt., Comanor, 
56la & 561b (NY, US). Puttalam District: Palavi Atta Villu, Cooray 6 Oct. 1969 
(NY); Puttalam, Trimen Aug. 1883 (PDA). Trincomalee District: Kantalai Tank, 
Mile 134 on Rd. A 6, Davidse 7518 (MO, NY). Mannar District: Aruvi Aru, Mile 
125 on Rd. A 4, T. Koyama et al. 13934B (NY); Palavi, Cooray in 1969 (PDA). 
Anuradhapura District: Anuradhapura, Trimen Oct, 1883 (PDA). Matale District: 
Ereua Tank, ca. 5 miles ESE of Dambulla, 650 ft. alt., Davidse 7391 (MO, NY, 
PDA, US). Colombo District: Colombo, Ferguson C.P. 3947 (K, PDA). Hamban- 
tota District: Ruhuna National Park, Attaville, Cooray 6 Oct. 1969 (NY). 


Distribution. Widely spread in Eurasia, from Mediterranean Region and East 
Africa through Asia Minor and India to southeastern Asia, Malesia and Australia. 


PYCREUS P. Beauvois 


1. Achenes smoothish to puncticulate with isodiametrical or hexagonal epidermal 


cells 


2. Culms few- to several-nodose below the middle, the lower part decumbent 
or obliquely ascending, branching and rooting at lower nodes; glumes 
furrowed on both sides of costa ..:...0..s..scsesevesceevee 39 P. sanguinolentus 


2. Culms not nodose above the base, erect from very base; glumes not 
furrowed 


Ceylon Cypereae 147 


3. Culms 20-90 cm tall; glumes acute to subobtuse at apex, never 
emarginate nor cuspidate 


4. Glumes orbicular-obovate, ca. 1 mm wide in half view, the margins 
very broadly whitish-hyaline, rounded to apex, spikelets 3 mm wide; 
culms 2—3 mm thick, mostly solitary ............ 40 P. puncticulatus 

4. Glumes ovate to lance-ovate, ca. 0.5 mm wide in half view, hardly 
or very narrowly hyaline on margins, acute at apex; spikelets ca. 
1.5 mm wide; culms 0.7-2 mm thick, as a rule tufted 


5. Lowest bract as long as to at most 2.5 times as long as the 
umbel, when longer than that, then inflorescence congested in 
a head; spikelets rusty brown to yellowish brown .................. 
- eee RUM Sidtnn -adteon pbinwng- Eh cate redaea, 41 P. polystachyos 


5. Lowest bract 3 to 5 times as long as the umbel; spikelets dark 
purplish-brown to chestnut-brown .................. 42 P. flavidus 


3. Culms 1-8 cm tall; glumes truncate to emarginate at apex, the midvein 
excurrent beyond the glume apex into a recurved mucro 


6. Glumes ovate, close together not exposing achenes; rays normally 
Porry tre veloped y. 660. Sete LS 43a P. pumilus ssp. pumilus 


6. Glumes oblong to lance-oblong, spaced, so that achenes exposed 
between glumes; rays well developed ......................cccccecee eee 
Mes See sone snvoXe eas enc <5 dtoh es 43b P. pumilus ssp. membranaceus 


38. Pycreus stramineus (Nees) C. B. Clarke, Fl. Brit. Ind. 6: 589, 1893; Alston 
in Trimen & Hook. f., Handb. Fl. Ceylon 6: 306, 1931. 


Cyperus stramineus Nees in Wight, Contrib. Bot. India 74, 1834; Trimen, 
Handb. FI. Ceylon 5: 19, 1900. Not of Desf. ex Link, 1820. Cyperus sub- 
stramineus Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 398, 1936; Kern, Fl. Males. 
Ser. 1, 7 (3): 653, 1974. 

Kurunegala District: Kurunegala, Trimen C.P. 3776 (PDA). 

Distribution. India, Ceylon, Indo-China and Malay Peninsula. 


Thus far known from Ceylon by a single collection only. This species closely 
resembles P. flavescens of much wider distribution, from which it differs primarily 
in its longer spikelets that are 8 to 35 mm long bearing many acute-tipped glumes 
in contrast to shorter spikelets (6 to 10 mm in length) with several obtuse 
glumes in the latter, . 


39. Pycreus sanguinolentus (Vahl) Nees [Linnaea 9: 283, 1835. Invalid combina- 
tion] ex C. B. Clarke, FI. Brit. India 6: 590, 1893; C. E. C. Fischer in Gamble, FI. 
Presid. Madras 9: 1627, 1931; Alston in Trimen & Hook. f., Handb. Fl. Ceylon 6: 
307, 1931. 

Cyperus sanguinolentus Vahl, Enum. Pl. 2: 351, 1806; Nees in Wight, 
Contrib. Bot. India 75, 1834; Thwaites, Enum. Pl. Zeyl. 342, 1864; Trimen & 
Hook. f., Handb. Fl. Ceylon 5: 20, 1900; Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 
385, 1936; Kern, Fl. Males. Ser. 1, 7 (3): 646, 1974. Cyperus eragrostis 
Vahl, Enum. Pl. 2: 322, 1806; Trimen, Syst. Cat. Flow. Pl. & Ferns Ceylon 99, 
1885. Not of Lamarck, 1791. 

Wilpattu National Park: Manikepola Uttu, 7. Koyama et al. 13460 (NY, 
PDA, US). Kandy District: Peradeniya, near water fall above University campus, 
Comanor 690 (NY): ca. 5 miles S.E. of Gampola, at Mile 18/6 on Gampola- 
Nuwara Eliya Rd., 720 m alt., Davidse et al. 7924 (MO, NY). Ratnapura District: 
11 miles E. of Diniyaya on Rd. A17, 750 m alt. Davidse 7586 (MO, NY). 
Amparai District: Senanaike Samudra, Padagoda, T. Koyama et al. 13983 (NY. 
PDA, US). 


Gardens’ Bulletin, Singapore — XXX (1977) 


148 


ne. 


G, lateral view of ache 


, habit. B & C, two views of prophyll. 
F, glume. 


ikelet. E, portion of rhachilla with two flowers. 


spi 
Scales = 1 mm. 


Fig. 7. Pycreus puncticulatus (Vahl) Nees. A 


D, 


Ceylon Cypereae 149 


Rather widely distributed in the Old World from northeastern Africa through 
Afghanistan, India and Indo-China northeastwards to China and Japan, eastwards 
to Malesia. 


40. Pycreus puncticulatus (Vahl) Nees, Fl. Brasil. 2 (1): 10, in note, 1842: C. B. 
Clarke, Fl. Brit. India 6: 593, 1893; C. E. C. Fischer in Gamble, Fl. Presid. Madras 
9: 1628, 1931; Alston in Trimen & Hook. f., Handb. Fl. Ceylon 6: 307, 1931. 
Fig. 7. 


Cyperus puncticulatus Vahl, Enum. Pl. 2: 348, 1806; Thwaites, Enum. PI. 
Zeyl. 342, 1864; Trimen & Hook. f., Handb. Fl. Ceylon 5: 21, 1900; Kiikenthal, 
Pflanzenr. 4 (20), 101 Heft: 362, 1936. Cyperus baccha Kunth, Enum. 
ee 2 BSS S357. Pycreus baccha (Kunth) Nees, [Linnaea 9: 283, 1835. 
Invalid combination] Fl. Brasil. 2 (1): 10, 1842. In note. 


Trincomalee District: Mavadichchenai, Alston 544 (PDA); Trincomalee, 
Glenie C.P. 3751 in part (PDA). Matale District: Dambulla, Thwaites C.P. 3751 
(K, PDA). Kurunegala District: Kurunegala, Ranasinghe 843 (PDA). Galle 
District: Haburagala, Amaratunga 2315 (PDA). Amparai District: E of Kunukala 
Kalapuwa Lagoon, ca. 4 miles S of Panama, sea level, T. Koyama et al. 14027 
(NY, PDA, US); Helawe Eliya, ca. 7 miles S of Panama, E of Helawe Lagoon, 
sea level, T. Koyama et al. 14025 (NY, PDA, US). Ruhuna National Park: 
Uraniyawala, ca. 1 mile W of Buttawa, Fosberg et al. 51018 (US); Palugaswela, 
Ecology Project Block I, Cooray 23 Mar. 1970 (NY). 


Distribution. Rather sporadically noted from southern India (Bombay, Mysore 
and Madras), Malay Peninsula and Cochinchina. 


In the genus Pycreus this species is well demarcated by its relatively loose, 
large umbel with broad, red-brownish spikelets, in which it resembles superficially 
Cyperus procerus, a stoloniferous perennial with trigonous achenes. 


41. Pycreus polystachyos (Rottboell) P. Beauvois, Fl. d;Oware 2: 48. ¢. 8&6, f. 2, 
1807; C. B. Clarke in Hook. f., Fl. Brit. India 6: 592, 1893: Alston in Trimen 
& Hook. f., Handb. Fl. Ceylon 6: 307, 1931. 


Cyperus polystachyos Rottboell, Descr. Icon. Rar. Nov. Pl. [llustr. 39, 
t. 11, f. 1, 1773; Nees in Wight, Contrib. Bot. India 75, 1834; Thwaites, Enum. 
Pl. Zeyl. 342, 1864; Trimen & Hook. f., Handb. Fl. Ceylon 5: 20, 1900; Kuken- 
thal, Pflanzenr. 4 (20), 101 Heft: 367, 1936; Kern, Fl. Males. Ser. 1, 7 (3): 649, 
1974. Cyperus paniculatus Rottboell, Descr. Icon. Rar. Nov. Pl. Illustr. 
40, 1773. Pycreus paniculatus (Rottb.) Nees [Linnaea 9: 283, 1835. Invalid 
combination] ex Alston in Trimen & Hook. f., Handb. Fl. Ceylon 6: 307, 1931. 
Cyperus polystachyos Rottb. var. Jaxiflorus Bentham, Fl. Austral. 7: 261, 
1878; Kiikenthal, Pflanzenr. 4 (20), Heft. 101: 370, 1936. Pycreus poly- 
stachyos P. Beauv. var. laxiflorus (Bentham) C. B. Clarke, Fl. Brit. India 6: 592, 
1893. 


Jaffna District: Jaffna, Ferguson C.P. 800 (PDA). Anuradhapura District: 
Anuradhapura, Alston 638 (PDA). Wilpattu National Park: Kuda Patessa, 
Wirawan et al. 1035 (US); Kollankanatta, Mueller-Dombois et al. 27 Apr. 1969 
(US); NE. of Kuruttu Pondi Villu, T. Koyama 7560 (NY). Trincomalee District: 
Trincomalee, Glenie C.P. 800 (PDA); Ottawady, valley of downstream of Mahaweli 
Ganga, ca. sea level, T. Koyama 13652 (NY, PDA, US); Mile 97 on Rd. A 15, 
Davidse 7560 (MO, NY). Polonnaruwa District: ca. 1 mile NE of Elahera along 
Amban Ganga, 400 ft. alt., Davidse 7357 (MO, NY). Batticaloa District: Batticaloa, 
Thwaites C.P. 800, March 1868 (K, PDA). Nuwara Eliya District: Ramboda Rd., 
near Ramboda, 3200 ft., Ballard 1136 (K, NY); Maskeliya, Wright 11 Nov. 1926 
(PDA). Matale District: Bata-Anduwa Gap, Sevastine, at marker 34/6, 1300 m 
alt., Tirvengadum et al. 17 (PDA). Kandy District: Hantane, Silva 296 (PDA); 
Peradeniya, Alston 280 (PDA): Gampola-Dolosbage Rd., Grupe 159 (PDA). 


150 Gardens’ Bulletin, Singapore — XXX (1977) 


Kurunegala District: Kurunegala, Thwaites C.P. 800 (PDA); Beddegama, Alston 
1397 (PDA). Colombo District: Muturajawela, Amaratunga, 18 Jan, 1971 (PDA). 
Galle District: Horawala, left bank of Bentota Ganga, Amaratunga 2495 (PDA), 
Haburagola, Amaratunga 2381 (PDA). Amparai District: Tandiadi Kalapu 
Lagoon, S of mile 215 on Road A4, T. Koyama et al. 14018 (NY, PDA, US); 
3 miles NW of Maha Oya, 50 m alt., Davidse et al. 9008 (MO, NY.); Inginiyagala 
National Park, between Baduluwela and ‘‘Westminster Abeey”’, 7. Koyama et al. 
13995 (PDA). Ruhuna National Park: Uraniya, Ecology Project Block I, Cooray 
22 Mar. 1970 (PDA, US). 


Distribution. Cosmopolitan, tropical and subtropical regions, The range is 
extending into warm-temperate regions in eastern Asia as far as to central Japan, 
and into Mediterranean Region. 


This species is extremely variable in the color of spikelets and in the 
inflorescences, which vary from a rather open umbel with spicately disposed spike- 
lets to a head through an intermediate state bearing fascicled spikelets on short 
rays. In Ceylon plants with open inflorescences are common. Although Alston (in 
Trimen & Hook. f., Handb. Fl. Ceylon 6: 307, 1932) listed P. ferrugineus as 
occurring in Ceylon without citation of specimens, I have so far been unable to 
see any documentation of this essentially African species from Ceylon and India. 
Pycreus ferrugineus distinctly differs from P. polystachyus at least in its spreading 
spikelets that are much broader than those of the latter at 2 to 3 vs. 1.25 to 
1.5 mm in width, and it can no way be confused with the latter. Hence, Alston’s 
note under P. ferrugineus mentioning that “This is doubtfully distinct from P. 
polystachyus,” and ‘‘Up to 5000 ft.; common” implies that his P. ferrugineus may 
constitute a misidentification for P. polystachyos itself. 


42. Pycreus flavidus (Retzius) T. Koyama, Journ. Jap. Bot. 51 (10): 313, 1976. 


Cyperus flavidus Retzius, Obs. Bot. 5: 13, 1788; Vahl, Enum. PI. 2: 334, 
1806; Kern, Fl. Males. Ser. 1, 7 (3): 648, 1974. [Cyperus globosus Allioni, 
Auctuar. Fl. Pedemont. 49, 1789; Trimen & Hook. f., Handb. Fl. Ceylon 5: 21, 
1900; Kikenthal, Pflanzenr. 4 (20), 101 Heft: 352, 1936. Not of Forskael, 1775.] 
Cyperus strictus Roxburgh, Fl. Ind. ed. 1, 1: 203, 1820. Cyperus 
capillaris Konig ex Roxburgh, l.c. 1: 198, 1820; Nees in Wight, Contrib. Bot. 
Ind. 76, 1834. Pycreus globosus (Allioni) Reichenbach, Fl. Germ. Excurs. 
140, 1830. Pycreus capillaris (Konig ex Roxb.) Nees [Linnaea 9: 283, 
1834. Invalid combination] ex C. B. Clarke, FI. Brit. India 6: 591, 1893. 
Pycreus strictus (Roxb.) Alston in Trimen & Hook. f., Handb. Fl. Ceylon 6: 307, 
1931, “Cyperus flavescens L.” sensu Thwaites, Enum. Pl. Zeyl. 342, 1864. 


Polonnaruwa District: ca. 1 mile N.E. of Amban Ganga, 400 ft. alt. Davidse 
7357A (MO). Kandy District: ca. 5 miles SE of Gampola at mile 18/6 on Road 
A5, 720 m alt., Davidse et al. 7925 (MO, NY); ca. 2 miles E, of Maddakele, slope 
N.W. of Krinckles, 1440 m alt., Davidse 8332 (MO); Pallekale, Alston 278 (PDA); 
Gordindihela, Willis, 27 Feb. 1906 (PDA); Upper Hantane Rd., above the University 
Campus, Comanor 317 (PDA, US). Kurunegala District: Kurunegala, Gardner 
C.P. 801 (PDA). Nuwara Eliya District: Nuwara Eliya, Trimen C.P. 801 (PDA), 
Amaratunga 230 & 1305 (PDA); Pattipola, Lazarides 7291 (PDA, US); between 
Hakgala and Nuwara Eliya, 5400 ft., Ballard 1269 (K, NY); Corbett’s Gap, 4000 
ft., Ballard 1050 (K, NY); Moon Plains, 6000 ft., Ballard 1217A (K, NY). Ratna- 
pura District: 11 miles E of Deniyaya at mile 62 on Rd. A17, 750 m alt., Davidse. 
7884 (MO, NY); Ratnapura, Thwaites C.P. 801 (K, PDA). Colombo District: 
Mount Lavinia, Trimen Oct. 1881 (PDA). Monaragala District: ca. 3 miles W. of 
Wellawaya at mile 135/10 on Rd. A4, 990 ft., Davidse 7742 (MO, NY). Ruhuna 
National Park: Kumbukkan Oya, Ecology Project Block II, Cooray 31 July 1969 
(NY, US). Hambantota District: 2 miles E of Katuwana, Lazarides 7350 
(PDA, US). , 


Ceylon Cypereae 151 


Widely distributed from Mediterranean Region and tropical Africa eastwards 
to Central and southern Asia, and northeastwards to Japan. 


This widespread species is variable particularly in the width and the color 
of spikelets. Ceylonese plants from up countries at altitudes of over 750 m generally 
possess strongly compressed, dark purplish-brown spikelets that are mostly less 
than 1.5 mm in width, a form resembling plants from Japan and temperate India. 
A few specimens collected at low altitudes such as Cooray 31 July 1969 and 
Davidse 7357 4A cited above bear less compressed, broader spikelets with stramineous- 
brown scales. 


43. Pycreus pumilus (L.) Nees [Linnaea 9: 283, 1935. Invalid combination] ex 
C. B. Clarke, Fl. Brit. India 6: 591, 1893, concerning the nomenclature, but 
excluding basionym; Domin, Bibl. Bot. Heft. 85: 417, 1915; C. E. C. Fischer in 
Gable, Fl. Madras 9: 1627, 1931. 


Cyperus pumilus L., Cent. Pl. 2: 6, 1756; Trimen & Hook. f., Handb. FI. 
Ceylon 5: 19, 1900; Kiikenthal, Pflanzenr. 4 (20), Heft. 101: 375, f. 44 A-E, 1936; 
Kern, Fl. Males. Ser. 1, 7 (3): 650, f. 66, 1974. Cyperus pluvinatus Nees 
& Meyen in Wight, Contrib. Bot. India 74, 1834; Thwaites, Enum. Pl. Zeyl. 342, 
1864. Pycreus nitens Nees, Nova Acta Nat. Cur. 19, Suppl. 1: 53, 1843; 
C. B. Clarke, Fl. Brit. India 6: 591, 1893. Pycreus pulvinatus (Nees & 
Meyen) Nees & Meyen ex Nees [Linnaea 9: 283, 1835, invalid combination, &] 
Nova Acta Nat. Cur. 19, Suppl. 1: 53, 1843. Dichostylis nitens (Nees) 
Palla, Bot. Jahrb. 10: 296, 1889. ““Pycreus patens (Vahl)” sensu Alston 
in Trimen & Hook. f., Handb. Fl. Ceylon 6: 306, 1931. Invalid combination. 


43a. Subsp. pumilus. 


Jaffna District: Jaffna Lagoon, north shore at ca. 10 miles SE of Navatkuli, 
T. Koyama et al. 14040 (NY, PDA, US); Jaffna, Gardner in 1846 (PDA). 
Anuradhapura District: Maradaukadewela, Trimen 21 Jan. 1896 (PDA); Wilpattu 
National Park, East Intermediate Zone, along Wildlife Dept. path at ca. 2 miles 
from barrier gate, T. Koyama & Jayasuria 13966 (NY, PDA, US). Polonnaruwa 
District: Tamankaduwa, Townsend 73/250 (K, US). Trincomalee District: Periya 
Kulam, ca. 7 miles NW of Trincomalee, 7. Koyama et al. 14064 (NY, PDA, US): 
Trincomalee, west side of Welcomble Hotel. Wheeler 1241] (PDA). Mannar 
District: Aruvi Aru, mile 125 on Rd. Al4, T. Koyama et al. 13931 (NY, PDA, 
US). Puttalam District: Chilaw, Trimen, Dec. 1880 (PDA). Kandy District: Pera- 
deniya, Thwaites C.P. 806 in part (PDA); Gannoruwa, Alston 298 (PDA); Hara- 
gama, Alston 3 Oct. 1926 (PDA). 


Distribution. From India through Indo-China and southern China eastwards 
to Malesia and Australia (rare in Queensland). 


Alston (l.c., 1931) applied Cyperus patens Vahl to the Ceylonese plants then 
passing as Cyperus pumilus, and proposed a combination, Pycreus patens (Vahl). 
Typical C. patens from Africa closely resembles subsp. membranaceus, and it may 
not be sufficiently different from the latter. However, C. patens without doubt does 
not represent the common phase of Ceylonese C. pumilus. 


43b. Subsp. membranaceus (Vahl) T. Koyama, stat. nov. 


Cyperus membranaceus Vahl, Enum. PI. 2: 330, 1806. Cyperus nitens 
Retz. var. membranaceus (Vahl) Bockeler, Linnaea 35: 484, 1868. Cyperus 
pumilus L. var. membranaceus (Vahl) Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 376, 
1935. Cyperus pumilus L. forma membranaceus (Vahl) C. B. Clarke, Journ. 
Linn. Soc. 21: 44, 1884. 


Hambantota District: Ruhuna National Park, between the Entrance and 
Ecology Project Plot R9, Cooray s.n., 11 Dec. 1969 (NY, US); Uraniya, Cooray, 
22 Mar. 1970 in part (NY). 


152 Gardens’ Bulletin, Singapore — XXX (1977) 


Distribution. India, Ceylon and Thailand; rather frequent in southern India, 
but rare in Thailand. 


Cooray’s record, cited above, makes a new entry of subsp. membranaceus to 
the flora of Ceylon. The difference between subsp. membranaceus and the typical 
phase exists mainly in the relatively minor details of floral glumes and achenes, In 
subsp. membranaceus the oblanceolate to oblong-obovate glumes are only ca. 1/3 mm 
wide in half view, being broadest much above the middle, and are considerably 
loosely disposed on the rhachilla exposing the greater portion of achenes between 
the glumes just as in Cyperus tenuispica, whereas in subsp. pumilus its broader 
elliptic to ovate-elliptic glumes are 1/2 to 3/5 mm wide in half view being broadest 
just below the middle, and are closely arranged, there being no space between two 
glumes, In general, the achenes of subsp. membranaceus are truly obovate and 
cannot be seen through the membranous glumes, while in subsp. pumilus its 
elliptic to lance-elliptic achenes are usually seen through the hyaline glumes. The 
spikelets of subsp. membranaceus are shorter and more laxly disposed on well 
elongated rays in contrast to the usually more congested nearly head-like inflores- 
cence in subsp. pumilus. Though minute, these differances are sufficiently consistent 
as far as I have examined materials, and hence I regard subsp. membranaceus a 
valid taxon. 


MARISCUS Gaertner 
1. Spikelets more or less flattened with acute edges, bearing 3 to many achenes; 
glumes folded with a conspicuous keel, laxly holding achene 


2. Plants perennial; culms more than 45O cm tall; leaves septate-nodulose; 
glumes without recurved awn 


3. Spikelets lanceolate, 2 mm wide, straw-coloured, disposed in a spike 
scaseh desde de cild abs dome RhRe ENE oy ke ey Uieed DRDEIS Mecreee ase ae ean 44 M. javanicus 


3. Spikelets linear, 1 mm wide, reddish-brown, densely congested in a 


globose head .£.0 27S eet te Se ee 45 M. compactus 
2. Plants annual; culms 4-20 cm tall; leaves not septate-nodulose; glumes with 
recurved awn, heance spikelets squarrose .................. 46 M. squarrosus 


1. Spikelets terete without conspicuous edges, bearing 1 or 2 achene(s) only; 
glumes involute without distinct keel, tightly surrounding achene 


4. Rhachilla slender, not at all spongy-thickened; spikelets 1- or 2-fruited 


5. Base of culms hardly enlarged or slightly globose with a corm-like 
enlargement; basal leaf sheaths purplish-brown 


6. Spikelets mostly 2-fruited, greenish to greenish-straw-coloured, 
patent to ascending at maturity, mostly 4-5 mm long; achenes 
2 mm long 


7. Spikes umbellate with elongated rays; spikelets spreading or 
the lower ones weakly reflexed at maturity ........................ 


7. Spikes subsessile, nearly capitate; spikelets at least the lower 
ones obliquely patent at maturity 


8. Rhizome short, not stoloniferous; leaves 2-5 mm wide; - 
umbel truly terminali(.9.... .VoOU yO a ee 
£6 SUR TT i ee 48a M. cyperinus ssp. cyperinus 


8. Rhizome emitting slender stolons; leaves as a rule 
0.7-1 mm wide; umbel quasi-lateral with the lowest bract 
continued down to the culmx..)..°2i0001. cai 


Ceylon Cypereae 153 


6. Spikelets 1-fruited, whitish-green, spreading at maturity, mostly 

2-3 mm long; achenes ca. 1.3 mm long ............ 49 M. paniceus 

5. Base of culm quasi-bulbose, i.e, with an ovoid to ovoid-oblong 
thickening clothed with leaf sheaths; basal leaf sheaths straw-brown. 

9. Rhizome emitting slender stolons; spikes sessile yet recognizable. 

a as a cp ncn boacne «a mpen = deci use 50 M. clarkei 

9. Rhizome loosely tufted without stolons; spikes congested in an 

ovoid or conical head, not readily recognizable ........................ 


I Ps tka ones ane psec Pend Ras antipins spies S66 = «areal 51 M. dubius 
4. Rhachilla spongy-thickened; achenes sunken in a depression of spongy 
rhachilla-internodes; spikelets 1-fruited ............... 52 M. pedunculatus 


44. Mariscus javanicus (Houttuyn) Merrill & Metcalfe, Lingnan Sci. Journ. 21: 
4, 1945. 


Cyperus javanicus Houttuyn, Nat. Hist. I], 13 Aanw. Pl. (1). ¢. 8&8 f. J, 
1782; Kern, Fl. Males. I, 7 (3): 635, f. 63, 1974. Cyperus pennatus Lamarck, 
Illustr. 1: 144, 1791; Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 476, f. 53 A-G, 1936. 
— Mariscus albescens Gaudichaud in Freyc., Boy. Bot. 415, 1826; C. B. 
Clarke, Fl. Brit. India 6: 623, 1893; Trimen & Hook. f., Handb. Fl. Ceylon §: 40, 
1900. ——- Mariscus pennatus (Lamarck) Domin, Bibl. Bot. Heft. 85: 440, 1915. 


Wilpattu National Park: Manikepola Uttu, 7. Koyama 13456 (NY, PDA, US). 
Anuradhapura District: Ritigala Strict Natural Reserve, approach to Weweltenna 
along southern slope, 700 ft. alt., Jayasuria 1294 (PDA); Anuradhapura, Trimen 
in 1881 (PDA). Trincomalee District: Mile 79 on Road A 4, Davidse 7558 (MO, 
NY). Batticaloa District: Batticaloa, Trimen C.P. 678 p.p. (PDA, K). Monaragala 
District: Crossing of Rd. A 2 and Kirindi Oya at Mile 183/2, ca. 15 miles S 
of Wellawaya, 400 ft. alt., Davidse 7760 (MO, NY, PDA, US). Ratnapura District: 
Raigam Korale, Thwaites C.P. 678 p.p. (PDA). Galle District: Bentota, Ballard 
1507 (K, NY). Hambantota District: N of Kataragama, Wirawan 644 (PDA); 
Ruhuna National Park: Block 3, Cooray 23 May 1968 (PDA, US). 


Distribution. Tropical Africa, Madagascar through Indian Subcontinent to 
southern China and the southern Ryukyus, as well as to Malesia, northern Australia 
and the Pacific Islands. 


45. Mariscus compactus (Retzius) Boldingh, Zakfi. Landb. Java 77, 1916; Druce, 
Rep. Bot. Exch. Club Brit. Isls. 1916: 634, 1917; Fischer in Gamble, Fl. Madras 
9: 1645, 1931. ————- Fig. 8 


Cyperus compactus Retzius, Osb. Bot. 5: 10, 1789; Kiikenthal, Pflanzenr. 4 
20), 101 Heft: 423, 1936; Kern, Fl. Males. I, 7 (3): 638, 1974. Cyperus 
dilutus Vahl, Enum. Pl. 2: 357, 1806. —— Mariscus microcephalus Pres]. Reliq. 
Haenk. 1: 182, 1828; C. B. Clarke, Fl. Brit. India 6: 624, 1893; Trimen & Hook. f., 
Handb. Fl. Ceylon 5: 40, 1900. —— Mariscus dilutus (Vahl) Nees in Wight, 
Contrib. Bot. India 90, 1834. 


Badulla District: ca. 8 miles E of Mahiyangana, 3.5 miles N of Mile 53 on 
Mihiyangana-Padiyatalawe Rd., T. Koyama et al. 13979 (NY, PDA, US). 

Distribution. India, southern continental China, Taiwan and Malesia. 

This species, easily recognizable by its characteristic globose spikes, occurs 
very sporadically in the low countries in Ceylon. I have not seen Gardner's 
collection from Kornegalle, which was cited by both Thwaites and Trimen as 


C.P. 815, and which seems to be the only other documentation of this species 
from Ceylon. 


46. Mariscus squarrosus (L.) C. B. Clarke, Fl. Brit. India 6: 623, 1893, concerning 
the basionym. 


154 Gardens’ Bulletin, Singapore — XXX (1977) 


Cyperus squarrosus L., Cent. Pl. 2: 6, 1756; Kern, Fl. Males. I, 7 (3): 631, 
1974. Cyperus aristatus Rottboell, Descr. Pl]. Rar. Progr. 22, 1772; C. B. 
Clarke, Fl. Brit. India 6: 606, 1893; Trimen & Hook. f., Handb. Fl. Ceylon 5: 24, 
1900; Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 502, f. 55 F -J, 1936. Mariscus 
aristatus (Rottboell) Chermezon, Bull. Soc. Bot. France 85: 366, 1938; Tang & 
Wang, FI. Reipubl. Pop. Sin. 11: 178, pl. 59, 1961. 


Jaffna District: north shore of Jaffna Lagoon, ca. 10 miles SE of Navatkuli, 
sea level, T. Koyama et al. 14037 (NY, PDA, US); Keerimalai to Point Pedro, 
Clayton 5205 (K). Wilpattu National Park: near Sadpuda Kallu, 2 miles E of 
Kattankandal Kulam, 30 m alt., Fosberg et al. 50830 (NY, US). Trincomalee 
District: Trincomalee, Glenie C.P. 966 ex p. (PDA). Puttalam District: Puttalam, 
Ferguson in 1881 (PDA). Polonnaruwa District: Polonnaruwa, Clayton 5116 (K, 
PDA); 13 miles E of Polonnaruwa on Trincomalee Rd., Clayton 5135 (K, PDA); 
ca. 1 mile E of Kolakanaweli, Mile 56/7 on Batticaloa Rd., T. Koyama 13575 
(NY). Mannar District: Aruvi Aru, Mile 125 on Rd. A 14, T. Koyama et al. 
13934 (NY, PDA, US). Anuradhapura District: Galpitigala, a few miles from 
Ritigala, Ballard 1451 (K, NY). Vavunia District: Palayanalankulan, Mile 120 on 
Rd, A 14, T. Koyama et al, 13938 (NY, PDA, US). Kandy District: Hantane, 
Thwaites C.P. 966 (K, PDA); Giragama, Amaratunga 359 (PDA). Hambantota 
District: Tissamaharama, Trimen Dec. 1882 (PDA); Rhuhuna National Park, 
Kohombagaswala, Block I, Cooray et al, 22 Jan. 1969 (US); Uraniya, Cooray 
22 Mar. 1970. 


Distribution. Widely distributed in tropical and subtropical regions of the 
world, extending also into temperate regions in the Americas, where the range 
covers from southern Canada south to Argentina and Chile. In Ceylon common on 
wet sandy soil in open grasslands. 


In the Linnaean Herbarium the specimen marked as “‘sguarrosus” in Linnaeus’ 
hand represents this taxon, while the second sheet with a single inflorescence is 
Cyperus maderaspatanus. Consequently, Cyperus squarrosus is the correct name 
of what has generally been called C. aristatus. Although I place this species in 
Mariscus following Chermezon and Wang & Tang (locis cit.), as was already 
pointed out by O’Neill (Rhodora 44: 47, 1942) and Kern (op. cit.) the caducity 
of the rhachillas of this species is not constant. In Ceylonese specimens I noticed 
in a few specimens that the glumes fall off apart from the rhachilla, which 
eventually falls off the rhachis. This species, which is characteristically intermediate 
between Cyperus and Mariscus, requires more detailed anatomical studies which 
might elucidate the systematic link between the two genera. 


47. Mariscus sumatrensis (Retzius) T. Koyama, comb. nova 


Scirpus cyperoides L., Mant. 2: 181, 1771. [Kyllinga umbellata Rott- 
boell, Descr. Icon. Rar. Nov. Pl. 15, 1773, excluding ¢. 4 f. 2. Illegitimate name.] 
Kyllinga sumatrensis Retzius, Obs. Bot. 4: 13, 1786. Kyllinga 
umbellata Rottb. var, sumatrensis (Retz.) Willdenow, Sp. Pl. 1: 258, 1797. 
[Mariscus umbellata (Rottb.) Vahl, Enum. 2: 376, 1806. Combination based on 
illegitimate name. ] Mariscus sieberianus Nees [Linnaea 9: 286, 1835, nomen 
nudum] ex C. B. Clarke, Fl. Brit. India 6: 622, 1893, & Illustr. Cyper. t. 23 f. 5-6, 
1909; Trimen & Hook. f., Handb. Fl. Ceylon 5: 42, 1900. Mariscus cyperoi- 
des (L.) Urban, Symb. Antill. 2 (1): 164, 1900. Not of Dietrich, 1833. 
Cyperus cyperoides (L.) O. Kuntze, Rev. Gen. Pl. 3 (2): 333, 1898; Kiikenthal, . 
Pflanzenr. 4 (20), 101 Heft: 514, 1936; Kern, Fl. Males. I, 7 (3): 642, 1974. 


Nuwara Eliya District: Hakgala, Alston 633 (PDA), Mueller-Dombois May 
1968 (PDA, US); Talawakele, Bond 17 Feb. 1944 (PDA). 
Distribution. Tropical Africa, tropical and subtropical Asia, Malesia, northern 


Australia; also introduced to the West Indies. In Ceylon rarely found in up 
countries. 


Ceylon Cypereae 


155 


Fig. 8. Mariscus compactus (Retzius) Boldingh. A, habit. B, orifice of a cladoprophyll 
at the base of umbel ray. C, spikelet. D, prophyll. E, lateral view of two glumes showing 
a base forming the wings on the rhachilla. F, glume. G, fruiting pistil. H, achene. Scales 
= | mm. 


156 Gardens’ Bulletin, Singapore — XXX (1977) 


Fig. 9. Mariscus clarkei (T. Cook) T. Koyama. A, habit. B & C, two views of spikelet. 
D. eewrone subtending a spikelet. E, prophyll. F, fruiting glume. G, empty glume. H, achene. 
es = 1 mm. 


Ceylon Cypereae 157 


When this taxon is placed in Mariscus, its earliest valid name, Scirpus 
cyperoides, cannot be transfered because of Dietrich’s homonym. Next earliest name, 
Kyllinga umbellata, is illegitimate since Rottboell cited Scirpus cyperoides in the 
synonymy. Kyllinga sumatrensis, therefore, needs to be transferred to Mariscus to 
form the correct name of this species under the genus. 


48. Mariscus cyperinus (Retzius) Vahl, Enum. Pl. 2: 377, 1806; C. B. Clarke, FI. 
Brit. India 6: 621, 1893, & Illustr. Cyper t 12 f. 3-4, 1909: Trimen & Hook. f., 
Handb. Fl. Ceylon 5: 42, 1900; C. E. C. Fischer in Gamble, Fl. Madras 9: 1644, 
1931. 


Kyllinga cyperina Retzius, Obs. Bot. 6: 21, 1791. Cyperus cyperinus 
(Retzius) Suringar, Het. Gesl. Cyper. Mal. Archip. 154, ¢. 6 f. 10, 1898; Kiikenthal, 
Pflanzenr. 4 (20), 101 Heft: 518, 1936; Kern, Fl. Maels. L 7 (3): 641, 1974. 


48a. Subsp. cyperinus. 

Ceylon: without definite locality, Trimen C.P. 816 (PDA). Kandy District: 
ca. 2 miles N of Hunnasgiriya, near Mile 23, 380 m alt., Davidse 8457 (MO, NY); 
Pusselawa, Alston 1170 (PDA); Peradeniya, Thwaites CP. 816 (K, PDA), Amara- 
tunga 643 (PDA), Alston 593 (PDA); Morakande Estate, Alston 594 (PDA); 
Gannoruwa, Alston 851 (PDA); Gampola, Alston 592 (PDA). Matale District: 
33/7 Midlands, 1200 m alt., Tirvengadum et al. 14 (US); Matale, Tirvengadum et al. 
16 (PDA). Nuwara Eliya District: Ramboda Pass, 3200 ft. alt.. Ballard 1142A 
(K, NY). Badulla District: Rawanaella Waterfall, ca. 2 miles SE of Ella, 750 m 
alt., Davidse et al. 8868 (NY, PDA, US). Colombo District: Dewalgama, Amara- 
tunga 1162 (PDA): Ekala, Alston 2389 (PDA). 


Distribution. From India to Malesia and northern Australia, also in Eastern 
Asia extending northwards to southern China and the Ryukyus. Open grassy places 
in hilly countries. 


48b. Subsp. laxatus (C. B. Clarke), T. Koyama, stat. nov. 


Cyperus umbellatus C. B. Clarke var. laxatus C. B. Clarke, Journ. Linn. Soc. 
21: 201, 1888. Cyperus cyperinus Suringar forma curvata Suringar, Het 
Gesl. Cyper. Mal. Archip. 156, 1898; Kern, Fl. Males. I, 7 (3): 624, 1974. 
Mariscus tenuifolius Schrader ex Nees, Fl. Brasil. 2 (1): 46, 1842: C. B. Clarke, 
Fl. Brit. India 6: 622, 1893; Trimen & Hook. f., Handb. Fi. Ceylon 5: 43, 1900; 
C. E. C. Fischer in Gamble, Fl. Madras 9: 1644, 1931. 


Kandy District: Hunnasgiriya, collector unknown 590 (PDA); Peradeniya, 
Alston 387 (PDA), Thwaites C.P. 817, ex p. (PDA); Maturata, Thwaites CP. 
817 ex p. (PD). Matara District: Weligama, Alston 1172 (PDA). Monaragala 
District: ca. 3 miles W of Wellawaya, near Mile 135/10, 990 ft. alt., Davidse 7741 
(MO, NY). Ruhuna National Park: Ecology Project Prot R19, Cooray 11 Dec. 
1969 (US). 

Distribution. India to Malesia; not as common as the typical phase. 

Subspecies /axatus differs from the typical phase in its narrow habit with long- 
creeping slender stolons and the few-spiked contracted umbels that tend to become 
pseudo-lateral being subtended by the lowest leafy bract continued down to the 
culm. As far as the Ceylonese plants are concerned this combination of characters 
appears to be rather constant, on which basis I regard this slender phase to be a 
subspecies of M. cyperinus. 


49. Mariscus paniceus (Rottboell) Vahl, Enum. Pl. 2: 373, 1806; C. B. Clarke, FI. 
Brit. India 6: 620, 1893, incl. var. roxburghianus, & Illustr. Cyper. t. 22 f. 1-2, 
1909; Trimen & Hook. f., Handb. Fl. Ceylon 5: 41, 1900, incl. var. roxburghianus. 


158 Gardens’ Bulletin, Singapore — XXX (1977) 


[Schaenoides paniceus Rottboell, Descr. Pl. Rar. Progr. 15, 1772, provisional 
name. | Kyllinga panicea Rottboell, Descr. Icon. Rar. Nov. Pl. 15, ¢. 4 f. J, 
1773. Cyperus paniceus (Rottboell) Bockeler, Linnaea 36: 381, 1870, in 
part, incl. basionym; Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 526, 1936; Kern, Fl. 
Males. I, 7 (3): 643, 1974. 

Anuradhapura District: Ritigala Nature Reserve, 2400 ft. alt., Jayasuria 1710 
(NY, PDA). Mannar District: Aruvi Aru, Mile 125 on Rd. A 4, T. Koyama et al. 
13928 (NY, PDA, US). Polonnaruwa District: SE of Giritale Wewa, 7 miles N.W. 
of Polonnaruwa, Fosberg & Ripley 51935 (US); Polonnaruwa, Ripley 339 (PDA); 
Galoya Reservoir, near spilway, 270 m alt., Commanor 568 (NY). Nuwara Eliya 
District: Ramboda Pass, 3200 ft., Ballard 1142A (K, NY); Hakgala Botanic 
Gardens, Clayton 5783 (K, PDA); between Pusselawa and Ramboda, at Mile 32/8 
on N. Eliya Rd., 1100 m alt. 7. Koyama & Herat 13613 (NY, PDA, US). Matale 
District: ca. 8 miles ESE of Dambulla, ca. 900 ft. alt., Davidse 7433 (MO, NY). 
Kandy District: Hantane, Thwaites C.P. 814 (K, PDA); Hewaheta, Thwaites C.P. 
2878 (PDA); Maragala, Alston 1629 (PDA); Haragama, Alston 306 (PDA); 
Aladeniya, Amaratunga 644 (PDA); between Alagalla and Balana Rd., Comanor 
440 (PDA, US). Kurunegala District: Kurunegala, Thwaites C.P. 2878 ex p. 
(PDA); Kurunegala Rock, Alston 699 (PDA); Wariyapola, Trimen Aug. 1883 
(PDA); Siyambalatenna, Alston 596 (PDA). Ratnapura District: ca. 13 miles 
N.E. of Deniyaya at Mile 64 on Rd. A 17, 1050 m alt., Davidse 7899 (MO, NY, 
PDA, US). Galle District: Galle, Alston 289 (PDA). Ruhuna National Park: 
Rugamtota, Cooray 16 Nov. 1969 (NY, US). 


While in Ceylon I was able to investigate the variation ranges of leaf width 
and the intensity of the umbels in this very common species. My conclusion 
is that var. roxburghianus should not be retained. It was claimed to differ from 
the typical phase in the relatively broader leaves and larger umbels. 


50. Mariscus clarkei (T. Cook) T. Koyama, Journ. Jap. Bot. 51 (10): 313, 1976. 
Fig. 9 

Cyperus clarkei T. Cook, Fl. Presid. Bombay 2: 873, 1908; Kikenthal, 
Pflanzenr. 4 (20), 101 Heft: 540, f. 58, 1936. [Mariscus pictus Nees in 
Wight, Contrib. Bot. India 90, 1834, in part excluding type.] [Mariscus 
bulbosus C. B. Clarke, Fl. Brit. India 6: 620, 1893; C. E. C. Fischer in Gamble, 
Fl. Madras 9: 1644, 1931. Not. of Steudel, 1855.] [Kyllinga bulbosa Vahl, 
Enum. Pl. 2: 376, 1806, invalid name mentioned in note; ROmer & Schultes, Syst. 
Veg. 2: 247, 1817, in note.] 


Monaragala District ca. 11 miles W of Tanamalwila, 125 m alt., Davidse & 
Sumithraarachchi 8812 (MO, NY, PDA, US). Hambantota District: Ruhuna 
National Park, Kohombagaswala, Cooray 23 Nov. 1969 (NY, PDA, US). 


Distribution, Confined to southern India and Ceylon. 


This species, newly found in Ceylon, rarely grows on wet shallow soil over 
rock outcrops. It is well demarcated by the peculiar bulb-shaped base, which 
emits a few slender stolons. The umbel of this species resembles that of M. 
ala and hence it is never confused with M. dubius, another bulbose Mariscus 
in Ceylon. 


ob 4 Maem dubius (Rottboell) Kiikenthal ex Fischer in Gamble, Fl. Madras 9: 


Cyperus dubius Rottboell, Descr. Icon. Rar. Pl. 20, t. 4 f. 5, 1773; Kiikenthal, 
Pflanzenr. 4 (20), 101 Heft: 563, 1936; Kern, Fl. Males. I, 7 (3): 643, 1974. 
Cyperus kyllingaeoides Vahl, Enum. Pl. 2: 312, 1806. Mariscus dregeanus 
Kunth, Enum. Pl. 2: 120, 1837; C. B. Clarke, Fl. Brit. India 6: 620, 1893, & 
ney Cyper. t. 2] f. 1-6, 1909; Trimen & Hook. f., Handb. Fl. Ceylon 5: 39, 


Ceylon Cypereae 159 


Anuradhapura District: Ritigala Natural Reserve, on summit rock, 2500 ft. 
alt., Jayasuria 905 (PDA). Polonnaruwa District: Lankatilaka, Silva 182 (PDA). 
Kurunegala District: Kurunegala, Thwaites C.P. 2942 (K, PDA) & C.P. 855 (K, 
PDA). Kandy District: Aladeniya, Amaratunga 642 (PDA); between Balane and 
Alagalla, 440 m alt. Comanor 1194 (NY, US). Badulla District: Rawanaella Water- 
fall, ca. 2 miles SE of Ella, 750 m alt., Davidse et al. 8866 (MO, NY, PDA, US): 
Lunugala, Trimen Jan. 1888 (PDA). Monaragala District: ca. 11 miles W of 
Tanamalwila, 125 m alt., Davidse et al. 8802 (MO, NY, US, PDA). Galle District: 
Galle, Gardner C.P. 855 (K, PDA). Ruhuna National Park: Bambowa, Cooray 
17 Nov. 1969 (NY, US); Rugamtota, Cooray 16 Nov. 1969 (NY, US). 


Distribution. Tropical Africa through India and Indo-China eastwards to 
Malesia. 


52. Mariscus pedunculatus (R. Brown) T. Koyama, comb. nov. 


Remirea maritima Aublet, Hist. Pl. Guian. Franc. 1: 45, ¢. 16, 1775. Not 
Mariscus maritimus Miquel, 1860, nor of C. B. Clarke, 1896. Remirea 
pedunculata R. Brown, Prodr. Fl. Nov. Holl. 236, 1810. Cyperus peduncu- 
latus (R. Brown) Kern, Act. Bot. Neerl. 7: 798, 1958. 


Jaffna District: Jaffna, Thwaites C.P. 3227 (K, PDA). Colombo District: 
Colombo, Ferguson C.P. 3227 (PDA); Mt. Lavinia, Trimen Oct. 1881. (PDA). 


Distribution. Southern Asia, Malesia and northern Australia; also in tropical 
America from eastern Venezuela through the Guianas south to Brazilian Parana 
Sand dunes and sandy shores of sea coast. 


The cyperoid morphology exhibited by this species was fully discussed by 
Kern (op. cit.), with whom I concur. When Cyperus sensu lato is divided into a 
few genera, this species properly belongs to Mariscus because of its continuous 
rhachilla jointed at its base. 


KYLLINGA Rottb. 
1. Inflorescence open, umbelliform with elongated rays ............... 53 K. hyalina 


1. Inflorescences congested in a head 
2. Glumes not winged 


3. Rhizome short; culms tufted 


4. Central spike cylindrical or oblong, 8-15 mm long, ca. 5 mm wide; 
spikelets broadly ovate, 2 mm long; culms slightly or hardly 
enlatmery at. Gase rt). £5.66 0265.5.0.5..5- 54 K. odorata ssp. cylindrica 


4. Central spike globose or ovoidal, 5-6 mm long, 3-4 mm across; 
spikelets oblong-elliptical, 1.7-2 mm long; culms bulbose-thickened 
(Ts RES \ 6 See Os ae epee eee a Oe 55 K. triceps 


3. Rhizome horizontally creeping or stoloniferous; culms distantly or close- 
ly arranged in one row along rhizome 


5. Culms 30-50 cm tall, close together 
Gem tier Practs 2 06:3 Ges eee. ieee eae 56 K. melanosperma 
i Fe Ode ee 57 K. polyphylla 
5. Culms as a rule 7-30 cm tall, more or less spaced ................... 
Te as J cemataele ihe inion 58 K. brevifolia 


SS ee eee pats. bt alti ea 59 K. nemoralis 


160 


’ Bulletin, Singapore — XXX (1977) 


Gardens 


Fy 
iA Z 


Afe a 
ti Lk 


co 


KY 44 
YT: 
“fp i 


/. F. 


Fig. 10. Kyllinga hyalina (Vahl) T. Koyama. A, habit. B, portion of rhachis with 
bracteoles and prophylls remaining after spikelets falling off. C, spikelet. D, prophyll. E, 
lateral view of two glumes. F, lateral view of achene. Scales = 1 mm, 


Ceylon Cypereae 161 


33: as “tage yep (Vahl) T. Koyama, Journ. Jap. Bot. 51 (10): 313, 1976. 
ig. 

Cyperus hyalinus Vahl, Enum. Pl. 2: 329, 1806; Trimen & Hook. f. Handb. 
Fl. Ceylon 5: 19, 1900; Kikenthal, Pflanzenr. 4 (20), 101 Heft: 498, 1936: Kern, 
Fl. Males. I, 7 (3): 655, t. 68, 1974. “Cyperus pumilus L.”’: sensu Nees in 
Wight, Contrib. Bot. Ind. 74, 1834. “Pycreus pumilus’’ C. B. Clarke, FI. 
Brit. Ind. 6: 591, 1893, concerning description. Pycreus hyalinus (Vahl) 
Domin, Bibl. Bot. Heft 85: 417, 1915, in obs. Queenslandiella mira Domin, 
Bibl. Bot. Heft 85: 416, 4 JJ f. 7-13, 1915. Mariscopsis suaveolens Cher- 
mezon, Bull. Mus. Paris 25: 60, 1919. Mariscopsis hyalinus (Vahl) Ballard, 
Kew Bull. 1932: 457, 1932. Queenslandiella hyalina (Vahl) Ballard in 
Hook., Icon. Pl. 33: t. 3208, 1933. 


Anuradhapura District: Anuradhapura, Trimen 18 Dec. 1881 (PDA). Kuru- 
negala District: Kurunegala, Trimen C.P. 3787 (K, PDA); Kuliyapitiya, Simpson 
9159 (PDA). Ruhuna National Park; near turn off to Andunoruwa Wewa, Cooray 
10 Dec, 1969 (NY, PDA, US); Ecology Project Block I, Plot R 13, Cooray 7 
Dec. 1969 (US). 


Distribution, Tropical East Africa, Mascarene Is., India, Indo-China, Malesia 
and northern Australia. Sporadically occurring in dry grasslands. 


When we split the genus Cyperus of broad meaning into several genera of 
smaller size, the systematic attribution of Cyperus hyalinus deserves special mention. 
Morphologically this species doubtlessly falls within the generic criterion of 
Kyllinga, from which it slightly deviates in having open inflorescences and in the 
spikelets of which nearly all the glumes are fruit-bearing. However, in the genus 
Kyllinga, umbel rays really develop as seen in K. transitoria for instance, and 
further where in all the cyperoid genera both open and head-like inflorescences 
occur without any clear border, thus showing that these conditions of inflorescences 
do not warrant generic segregation. Similarly the number of fruit-bearing glumes 
in a spikelets also does not serve as a distinguishing character between genera. In 
Mariscus, the genus most closely rێlated to Kyllinga, both multi- and few-flowered 
spikelets coexist (cfr. M. cyperinus vs. M. javanicus, for example). It is therefore 
permissible to include Cyperus hyalinus in Kyllinga in spite of the multi-flowered 
spikelets. In reality Cyperus hyalinus strikingly resembles Kyllinga ssp. not only 
in its bilaterally flattened oblong-elliptic achenes but also in its thinly membranous 
glumes with several prominent veins on both sides of the midrib. 


54. Kyllinga odorata Vahl, Enum. Pl. 2: 382, 1906. 


This species, occurring in tropical America and Africa, is represented in 
Ceylon by: 


subsp. cylindrica (Nees ex Wight) T. Koyama, comb. nov. 


Kyllinga cylindrica Nees ex Wight, Contrib. Bot. Ind. 91, 1834; C. B. Clarke, 
Fl. Brit. Ind. 6: 588, 1893, incl. var. subtriceps Nees; Trimen & Hook. f., Handb. 
Fl. Ceylon 5: 44, 1900. Kyllinga odorata Vahl var. cylindrica (Nees) 
Kiikenthal ex Merrill, Journ. Str. Branch Roy. As. Soc. 76: 80, 1917. 
Cyperus sesquiflorus (Torrey) Mattfeld & Kiikenthal var. cylindricus (Nees) 
Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 593, 1936. Cyperus sesquiflorus 
Mattfeld & Kiikenthal subsp. cylindricus (Nees) T. Koyama, Bot. Mag. Tokyo 
83: 187, 1970. Cyperus sesquiflorus Mattfeld & Kiikenthal var. subtriceps 
(Nees) T. Koyama, Quart. Journ. Taiwan Mus. 14: 191, 1961; Kern, FI. Males. 
I, 7 (3): 659, 1974. 


Kandy District: Pusselawa, Alston 19 Sept. 1926 (PDA). Badulla District: 
Haldummulla, Thwaites C.P. 3754 (K, PDA). Nuwara Eliya District: Ohiya, 
Mueller-Dombois 18 May 1968 (PDA, US); Hakgala, Simpson 9041 (PDA); 


162 Gardens’ Bulletin, Singapore — XXX (1977) 


Ramboda, along N. Eliya Rd., 1180 m alt., T. Koyama & Herat 13619 (NY, PDA, 
US). Hambantota District: Ruhuna National Park, Cooray 30 November 1967 
(PDA, US). 


Distribution. Tropical Africa, tropical and subtropical Asia. 


The morphological differences between subsp. cylindrica and the typical phase 
were discussed in my previous paper (Koyama, op. cit. 1970). 


55. Kyllinga triceps Rottboell, Descr. Icon. 14, t. 4 f. 6, 1773; C. B. Clarke, FI. 
Brit. Ind. 6: 587, 1893; Trimen & Hook. f., Handb. Fl. Ceylon 5: 45, 1900. 


[Schaenoides triceps Rottboell, Descr. Pl. Rar. Progr. 15, 1772, invalid name.] 
Cyperus triceps (Rottb.) Endlicher, Cat. Hort. Acad. Vindb. 1: 94, 1842; 
Kikenthal, Pflanzenr. 4 (20), 101 Heft: 578, 1936; Kern, Fl. Males. I, 7 (3): 659, 
1974. 


Mannar District: Aruvi Aru, Mile 125 on Rd. A 14, T. Koyama et al. 13932 
(NY, PDA, US). Polonnaruwa District: Polonnaruwa Sacred Area, Sect. 3A, 
61 m alt., Ripley 334 (US); Polonnaruwa-Trincomalee Rd., Clayton 5136 (K, 
PDA). Matale District: 33/7 Midlands, Tilyengadum et al. 12 (US). Puttalam 
District: Anamaduwa, Trimen Aug, 1883 (PDA); Puttalam, Brodie C.P. 3234 in 
part (PDA). Colombo District: Colombo, Thwaites C.P. 3234 (K, PDA). Mona- 
ragala District: ca. 11 miles W of Tanamalwila, 125 m alt., Davidse et al. 8810 
(MO, NY, PDA). Ruhuna National Park: Komawa Wewa, Cooray 16 Nov. 1969 
(PDA, US); Gonalabbe, Cooray 24 Nov. 1969 (NY, PDA); Yala, Andunoruwa 
Wewa, Comanor 673 (NY, US). 


Distribution. From tropical Africa through India and Indo-China north- 
eastwards to southern China, and eastwards to Malesia and northern Australia. 


56. Kyllinga melanosperma Nees in Wight, Contrib. Bot. India 91, 1834; C. B. 
Clarke, Fl. Brit. India 6: 588, 1893; Trimen & Hook. f., Handb. Fl. Ceylon 5: 45, 
1900. 


Cyperus melanospermus (Nees) Suringar, Het. Gesl. Cyperus Mal. Archip. 
50, t. 2 f. 8, 1898; Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 583, 1936; Kern, FI. 
Males. I, 7 (3): 655, 1974. 


Kandy District: Peradeniya, Lower Hantana Rd., near the waterfall, 535 m 
alt., Comanor 692 (NY, US); between Pusselawa and Ramboda, Comanor 950 
(PDA); between Laxapana and Maskeliya Rd., Kostermanns 24079 (PDA); 
Ambagamuwa, Amaratunga 2014 (PDA), between Udahentenna and Nawalapitiya, 
Amaratunga 909 (PDA); Aladeniya, Amaratunga 1643 (PDA). Nuwara Eliya 
District: Hakgala, Wheeler 12371 (PDA); Nuwara Eliya, Thwaites C.P. 818 (K, 
PDA); Horton Plains, forest opposite to the Horton Plains Farm, along Pattipola 
Rd., 7200 ft. alt., 7. Koyama 13506 (NY, PDA, US); base of Hakgala Mtn. 1870 m 
alt., Davidse et al. 7981 (MO, NY, PDA, US). Badulla District: ca. 3 miles W of 
Koslanda, along Rd. A 4, ca. 900 m alt., T. Koyama et al. 14033 (NY, PDA, US). 
Colombo District: Wattala, Comanor 1153 (PDA, US); Danowita; Amaratunga 
193 (PDA); Kimbulapitiya, Amaratunga 2157 (PDA). Ratnapura District: 11 
miles E of Diniyaya, near Mile 62 on Rd. A 17, 750 m alt., Davidse 7875 (MO, 


NY, PDA, US). Galle District: Corbett’s Gap. 4500 ft. alt., Ballard 1052 (K, NY). 


__ Distribution. Tropical and subtropical Africa, southern Asia, Fiji and Malane- 
sia. In Ceylon common on the grassy beds of montane forests. 
This species, well characterized by its relatively thick leafless culms and 


large heads, can be easily located in the field because of its strong odor of 
essential oil, which is contained in the thick rhizome. 


ee ee 


Ceylon Cypereae 3 163 


57. Kyllinga polyphylla Willdenow ex Kunth, Enum. Pl. 2: 134, 1937. 


Kyllinga aromatica Ridley, Trans. Linn. Soc. II, Bot. 2: 146, 1884. 
Cyperus aromaticus (Ridley) Mattfeld & Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 
581, 1936; T. Koyama, Bot. Mag. Tokyo 83: 186, 1970; Kern, Fl. Males. I, 7 (3): 
656, 1974. 


Nuwara Eliya District: between Pussellawa and Ramboda, at Mile 28/10 along 
Kandy-N. Eliya Rd., 950 m alt., Comanor 328 (NY, PDA, US). 


Distribution, E. Africa. Also eoHEeS to Singapore, Ceylon, Solomon Islands, 
Samoa and Fiji. 


This African species can be separated from the closely allied indigenous 
species, K. melanosperma, by its head subtended by 6 to 8 leafy bracts in contrast 
to 2 or 3 in the latter. In Ceylon it is thus far known only by the collection cited 
above, and is apparently of rather recent introduction. 


58. Kyllinga brevifolia Rottboell, Descr. Icon. Rar. Nov. Pl. 13, t. 4 f. 3, 1773; 
C. B. Clarke, Fl. Brit. India 6: 588, 1893, & Illustr. Cyp. ¢. J f. 1-4, 1909; Trimen 
& Hook. f., Handb. Fl. Ceylon 5: 45, 1900. 


[Schaenoides brevifolius Rottb., Descr. Pl. Rar. Progr. 15, 1772, invalid 
name. |] Cyperus brevifolius (Rottb.) Hasskarl, Cat. Hort. Bogor. 24, 1884; 
Kiikenthal, Pflanzenr. 4 (20), 101 Heft: 600, 1936; Kern, Fl. Males. I, 7 (3): 656, 
f. 70, 1974. 


Wilpattu National Park: Kali Villu, Wirawan et al. 990 (US). Mannar 
District: Aruvi Aru, Mile 125 on Rd. A 4, T. Koyama et al. 13926 (NY, PDA, 
US). Kandy District: Peradeniya, Upper Hantana Rd., 550 m alt., Comanor 315 
(NY, US); Ambewela, Mueller-Dombois 12 Jan. 1968 (PDA, US). Nuwara Eliya 
District: Kande Ela Reservoir, Mueller-Dombois 18 May 1968 (PDA). Colombo 
District: Dompe, Amaratunga 2182 (PDA); Ranmuthugal, Amaratunga 2221 
(PDA). Galle District: Galle, Trimen C.P. 3755 (K, PDA). 


Distribution. Pantropic; in Asia extending northwards into the warm regions 
of continental China and the temperate region of Japan. 


This cosmopolitan weedy sedge is rather poorly documented in Ceylon. Since 
it frequently grows in and around rice fields, it is quite possible that this sedge 
extended its range with the introduction of rice as assumed for Cyperus iria and 
Fimbristylis miliacea. 


59. Kyllinga nemoralis Gg. R. & G. Forster) Dandy ex Hutchinson & Dalziel, FI. 
W. Trop. Africa 2: 486, in Key, & 487, 1936. 


[Kyllinga monocephala Rottboell, Descr. Icon. Rar. Nov. Pl. 13, t. 4 f. 4, 
1773, invalid name; C. B. Clarke, Fl. Brit. India 6: 588, 1893 & Illustr. Cyper. 
t, 2 f. 1-2, 1909; Trimen & Hook. f., Handb. Fl. Ceylon 5: 44, 1900.] Cyperus 
kyllingia Endlicher, Cat. Hort. Acad. Vindb. 1: 94, 1842; Kikenthal, Pflanzenr. 
4 (20), 101 Heft: 606, f. 64, 1936; Kern, Fl. Males. I, 7 (3): 659, 1974. 
Thryocephalon nemoralis J. R. & G. Forster, Char. Gen. Pl. 130, 1776. 


Trincomalee District: Mutur, Wheeler 12425 (PDA). Kandy District: 
Roseneath, 2101 ft. alt., Ballard 1014 (K, NY). Monaragala District: ca. 3 miles 
W of Wellawaya at Mile 135/10 on Rd. A 4, 990 ft. alt. Davidse 7728 & 7732 
(MO, NY, PDA, US). Ruhuna National Park: Rugantota, Block I, Cooray 16 
Nov. 1969 (NY, US); Ecology Project Block I, Rugantota on Minik Ganga, 
Fosberg et al. 50191 (NY, US). 


Distribution. Pantropic, relatively scarce in tropical America. 

It is unfortunate that to this common species, which has long been known as 
Kyllinga monocephala, must be assigned an unfamiliar epithet as its correct name. 
Rottboell’s Kyllinga monocephala is invalid as the name was published with an 
earlier legitimate name, Schoenus coloratus L., cited in the synonymy. 


164 Gardens’ Bulletin, Singapore — XXX (1977) 
Literature Cited outside Taxonomic Enumerations 


Kern, J. H. 1974. Cyperaceae. Flora Malesiana Ser. I, 7 (3): 435-753. 


Koyama, T. 1970. Beitrage zur Cyperaceenflora von Ceylon. Bot. Mag. Tokyo 
83: 183-192. 


1974. Additional notes on Ceylonese Cyperaceae. Bull. National 
Sci. Mus. Tokyo 17: 65-73. 


Kiikenthal, G. 1935-36. Cyperaceae-Scirpoideae-Cypereae. Pflanzenr. 4 (20), 
101 Heft: 1-671. 


Trimen, H. & J. D. Hooker. 1900. A Handbook to the Flora of Ceylon 5: 1-477. 


The Limestone Hill Flora of Malaya I 


by 
S. C. CHIN 
University of Malaya, Kuala Lumpur 
Malaysia 
CONTENTS 
‘PAGE 
Abstract : - - . - - : - - - 165 
Preface and Acknowledgements’ - - - - > - : 166 
SECTION | — GENERAL 
1. Introduction’ - : - - - - - - 167 
2. Geology, origin and distribution of the limestone hills - - 172 
3. Vegetation - - - ‘ = i > , 174 
4. Phytography and size of the flora - . . - : 182 
5. Affinities of the floristic components to limestone’ - - - 189 
SECTION 2— The Flora 
Pteridophytes : re : * - - - 198 
Gymnosperms - - - - - - - - 218 
Abstract 


Limestone habitats, chiefly karst towers, in the Malayan Peninsula, support a rich flora 
of about 1216 species of vascular plants, in 582 genera and 124 families. Phanerogam 
families total 119, representing 61.6% of the total number of phanerogam families recorded 
for the whole Malayan flora; only 72 families are not represented on limestone, and these 
are mostly aquatics or small rare groups. Specific endemism among the limestone plants 
is 21.4% (261 species), and of these 10.7% (130 species) are found only on limestone. There 
are 335 species “characteristic” of the limestone flora, and 254 of these (20.8%) are restricted 
to limestone. 


The limestone vegetation is described and classified into “types” and secondary vegetation 
and succession is discussed. Pioneer species on limestone include those found on other 
disturbed terrestrial habitats in Malaya. Some plants found on limestone are found elsewhere 
in Malaya only at significantly higher elevations. A discussion of the geological origin and 
distribution of the Malayan limestone areas is also included. 


165 


166 Gardens’ Bulletin, Singapore — XXX (1977) 


PREFACE AND ACKNOWLEDGEMENTS 


This publication is based on the dissertation accepted for the Master of 
Science Degree in 1973 by the School of Biological Sciences, University of Malaya. 
I am grateful to Professor van Steenis, external examiner, for his critical comments 
and helpful suggestions. 

My work was supervised by Dr. B. C. Stone who introduced me to this topic, 
provided an unfailing source of guidance, and who also permitted me free access 
to his most remarkable collection of botanical literature which has been an 
inspiring and invaluable source of reference. I am also grateful for his encourage- 
ment to publish this work. 


The Department of Botany, University of Singapore allowed me the use of 
their herbarium and Dr. H. Keng helped with the identification of several Labiatae. 
He also generously provided me with the records of the collection made by the 
UNESCO 1962 limestone expedition to Ulu Kelantan. These consisted of a map, 
several note books and a set of duplicate labels to the collection. 


The Director of the Botanic Gardens, Singapore, gave me permission to work 
and the staff helped me in the herbarium on several occasions. 

Dr. T. C. Whitmore and subsequently Dr. Francis Ng allowed me to use 
freely the facilities of the herbarium at the Forest Research Institute, Kepong, and 
manuscripts of their “Tree Flora of Malaya’, vol. 2 then in press. Dr. Whitmore 
also helped with the identification of some Garcinias, Euphorbiaceae and Palms; 
Dr. Ng with some Ebenaceae and other members of the staff with some general 
collections. 

Dr. K. U. Kramer of the State University, Utrecht, Netherlands, identified 
several fern specimens and Dr. T. Shimizu of Shinshu University, Japan, provided 
literature and helped with the identification of a specimen of Impatiens. 

With Dr. P. R. Wycherley, formerly of the Rubber Research Institute, Kuala 
Lumpur, I had interesting discussions and he also introduced me to the Johore 
limestone. 

Mr. W. Swinson, Project Manager for the South East Johore Project, then 
engaged in a Master Plan Study of South East and Central Johore for the Govern- 
ment of Malaysia and State Government of Johore and his staff provided maps, 
transport, guides and equipment for two trips into the Gunong Sumalayang 
limestone outcrops in Johore. Mr. Swinson also provided unpublished data on the 
locality. 7 | 

Dato Haji Wan Hassan bin Abdul Halim, the State Forest Officer of Johore 
gave permission to enter and botanize in the Johore limestone area. 

Incik Mahmud bin Sidek and Incik Badaruddin, field investigator and 
herbarium assistant respectively, assisted in several excursions. Friends, particularly 
J. Boey, provided help and interesting company on numerous trips to the limestone. 

To all these helpful persons. J express my most grateful thanks. 


I must also state my appreciation for the comments of Dr Chang Kiaw Lan 
editor. | | 

Most of this work was carried out during the tenure of a tutorship in the 
School of Biological Sciencés, Universiti Malaya. 


Malayan Limestone Flora 167 


SECTION 1— General 
I. INTRODUCTION | 


Background to the flora 


The limestone landscape in Malaya is very distinctive. Typically the hills are 
tower-like, rising from the surrounding scenery, with sheer rock walls and often 
jagged summits. These ‘tower karst’ formations occur as isolated crags or are 
grouped together into small massifs — all products of erosion of calcareous rock. 
Erosion too is responsible for the presence of caves that often characterize limestone 
hills. These caves have, since prehistoric times, often been used as human dwellings, 
and as much as the vegetation on the hills provide clues to the history and evolution 
of the Malayan flora, provide evidence for an insight into prehistoric Malayan 
culture. 

Most of the past work on the Malayan limestone concerned the geology, 
origin and age of the rock, e.g. Scrivenor (1931), Paton (1961, 1964), Gobbett 
(1965) and Hutchison (1963, 1968). Other investigators have concerned them- 
selves with cave archaeology, e.g. Evans (1920), Tweedie (1940) and Peacock 
(1965), or cave ecology and fauna, e.g. Bullock (1963, 1965), Dunn (1965), 
McClure (1965) and Tweedie (1947). 

The only separate work on the flora is that of Henderson (1939). This 
contains an introduction giving the collecting localities, with notes on the vegetation 
and the characteristics of the flora. The localities are introduced state by state; the 
chief hills are named and the major collectors mentioned. There is however, 
no map of the distribution of the hills mentioned. The vegetation is described; 
there is a division of the hills into two main groups, the ‘wet’ and ‘dry’ hills. The 
frequency of the occurrence of the plant families (restricted to the spermatophytes) 
is briefly summarized. 

A total of about 745 species is recorded from the limestone by Henderson. 
The species are presented in a check-list with short distributional notes; the 
arrangement of the families follows Ridley (1922-25), while the genera and 
species are arranged in alphabetical order. Of the 745 species recorded about 195 
(26%) in the Peninsula are known only from limestone. The species endemic and 
confined to the limestone number about 130. 


Apart from this study, species distribution on limestone is sometimes men- 
tioned in floristic or monographic works on the local flora. Thus this well known 
habitat is, botanically, relatively unworked. 


The history of botanical exploration on the Malayan limestone hills began in 
August 1880 when Kunstler, collector for Sir George King, made a collecting trip 
to the Gopeng limestone in Perak (the exact locality is unknown). Other early 
collectors (before 1900) include Curtis, Fox, Kelsall, Ridley and Wooldridge. Up 
to the present day more than 60 collectors have left their mark on this history. 
Their efforts have accumulated about 4,500 numbers (an estimate based on my 
having seen about 4000 numbers and assuming that I have not seen about 10% 
of the total); of this the writer contributed 1550 numbers. 


The total area of limestone in Malaya, considering the average height to be 
243.8 m (800 ft), is about 260 sq. km. (an estimate based on Scrivenor, in Burkill, 
1935). Therefore per unit area the limestone may seem to have been very intensively 
botanised, with about 1962 numbers per 100 sq. km. (compared with the average 
of 175 numbers per sq. km. for the whole of Malaya, Steenis-Kruseman, 1973). 
These figures are, however, deceptive and misleading when the richness of the 
limestone flora is taken into account. The total number of collections from the 
Malay Peninsula is about 232,000 (a rough estimate based on Steenis-Kruseman, 
1973) and the total species present in Malaya is estimated to be between 
8000-8500 (Keng, 1970). This total number of species is an estimation only and 
is more than the number of species presently known. Thus this shows that on the 


168 Gardens’ Bulletin, Singapore — XXX (1977) 


average, each species is represented by 26-29 numbers, On the other hand, 1216 
species are recorded from the limestone (present study) which are represented by 
about 4500 numbers, thus giving an average of about 4 numbers per species. Even 
if only the characteristic species (with affinities for the limestone habitat) were 
considered (they number 335 and the total numbers collected would also drop 
far below 4500) and the total numbers collected be retained at 4500, the average 
number of specimens per species will be 13, which is still low. 


Besides low total numbers collected with respect to the extremely rich flora, 
many of the limestone species have a very local distribution which intensifies the 
problem of insufficient collection. Though most of the limestone localities have 
been visited by botanists, the total number of hills actually climbed and collected 
on is only about 50, and the number of islands (in Langkawi) visited is about 25 
(the writer has visited 25 hills and 12 islands, some repeatedly). There are no 
estimates of the total number of limestone hills in Malaya, but they probably 
number well over 200, and Langkawi consists of 99 islands (though not all are 
limestone, a large percentage are). Thus we have a situation where many hills, 
islands and localities have been repeatedly visited while others have not been 
botanised at all. The “‘popular” localities include Bukit Takun, Gunong Rapat, 
Gua Musang, Pulau Dayang Bunting (around the lake), Pulau Jerkom and Pulau 
Timun. However, this is not to say that these relatively frequently visited spots are 
sufficiently botanised. For instance, recently I discovered a clump of Oncosperma 
horridum on the summit of Gua Batu (which rates high on the list of popular 
localities), a species which has not been recorded on the limestone before. Con- 
sidering that Gua Batu is less than 2.59 sq. km (1 sq. mile), in area, this clearly 
shows that the many rugged faces of a limestone hill can conceal even the most 
conspicuous plant. 


Therefore in relation to its rich flora, extremely varied habitat and the fact 
of uneven local exploration, the limestone vegetation is probably the least botanised, 
and hence botanically known, of all the vegetation types in Malaya. 


The collectors who have contributed to our knowledge of the limestone flora 
are listed below with a brief mention of collection dates and localities. 


Collectors 


Allen, B.E.G.M. Dec. 15th. 1950; Gunong Keriang (Kedah), a specimen of Chirita viola 
Ridl. 1958-1960; widely in Perak, visited Gunong Idong, Gunong Kanthan, Rotan Segar. 
Almost 100 numbers. 

Allen, E.F. May 1946; collected on ‘Kinta limestone’. April 1955-Jan. 1957; collected on 
several occasions from Gua Batu and Bukit Takun in Selangor. 

Alphonso, A.G. & A. Samsuri. Nov.—Dec. 1960; several numbers from Langkawi (Pulau 
Bumbon Besar and Bumbon Kechil). 

Batten Pooll, A.LH. A poorly labelled Dendrocalamus elegans (Ridl.) Holtt. from the Langkawi 
limestone. 

Bra Oct. 29th-Nov, 3rd, 1929; miscellaneous, about 80 numbers from Gunong Baling, 

edah. | 

Boey, H.Y. July 1970 — Sept 1971; collected from Bukit Takun on four occasions and once 
from Bukit Anak Takun, Selangor. August 1971; collected from Gua Musang, Gua Batu 
Boh, Batu Neng and Batu Tapah (all in Kelantan) and Gua Layang, Pahang. November 
1971; visited Langkawi and collected from a number of limestone islands. About 350 
numbers in all. 

Burkill, ILH. 1916-1920; collected from Gua Batu, Selangor and around Ipoh, Perak. Some- 
times with Haniff. 

Burtt, B.L. & Woods, P.J.B.; Langkawi, at Sungei Kisap and Pulau Dayang Bunting; Perlis 
at Bukit Chupeng and Bukit Bintang; Perak, around Ipoh. About 146 numbers in all. 

Carr, C.E. 1928-1930; visited Kota Glanggi, Gunong Senyum, Gua Tipus and Tembeling in 
Pahang. Collected mainly orchids; some ferns also. | 

Chan, Y.C. July 1970; collected from around Lenggong, Perak. Several assorted numbers. 

Chew, W.L. April-May 1957; collected on Bukit Hantu, Bukit Kalong, Gunong Baling and 
around Dayang Bunting, Langkawi; all in Kedah. In the same period also visited Bukit 
Manek and Kaki Bukit in Perlis. 


4 

. 

b 
~ 
a 


Malayan Limestone Flora 169 


Cockburn, P.F. June 10th 1968; some specimens from Batu Biwa in Trengganu. 


Corner, E.J.H. Nov. 16th—25th, 1929 (with Henderson); collected on Bukit Lagi, Perlis. 
1936-1937; visited Ipoh and Bukit Takun. Nov. 13th-22nd 1941 (sometimes with Nauen); 
collected from all over Langkawi, resulting in some 200 numbers. 25th Nov. 1941 (with 
Nauen); collected on Gunong Baling, Kedah. 


Curtis, C. 1888-1902; visited Langkawi (Kedah), Gua Batu in Selangor, Kinta district, around 
Ipoh and Sungei Siput in Perak. Most of the specimens are very poorly labelled. 

Durant, C.L. Oct. 30th 1940; a specimen of Maxburretia rupicola (Ridl.) Fur. from Bukit 
Takun (Selangor). 

Dransfield, J. July 3lst 1968; several orchids from Bukit Anak Takun (Selangor), 

Enoch, I. Sept. 25th 1954; a specimen of Impatiens mirabilis Hk.f. from Pahang. 

Evans, G.B. Early 1966; Ipoh Temple limestone (Perak) and Bukit Takun (Selangor); 
several ferns. 

Everett, B. July 1970; collected from around Lenggong, Perak. Eighteen numbers. 

Fox, W. 1899-1904; several plants from Langkawi (Kedah) and around Ipoh (Perak). 

Furtado. C.X. June 4th 1937; several palms and two aroids from Gunong Baling, Kedah. 


Haniff, Mohamed. 1900-1921; collected occasionally from limestone, visited Bukit Wang, 
Gunong Keriang and Langkawi (Kedah), and Gunong Pondok (Perak). After 1918 
usually with Nur. See also Burkill. 

Hashim, Ja’afar. Collected a Wikstroemia indica (L.) C.A. Mey from Langkawi. 

Henderson, M.R. 1923-1935; visited most of the limestone localities. Langkawi, usually on 
both sides of Selat Panchor (straits), and around Pulau Dayang Bunting. Gua Lambok, 
Gua Musang, Gua Panjang and Gua Teja (Kelantan). Bukit Cheras, Bukit Chintamani, 
Bukit Serdam, Gua Tipus, Gunong Senyum and Kota Glanggi (Pahang). Gua Lanno, 
Gunong Pondok, Ipoh, Lenggong and Pulai (Perak). Besih Hangat, Bukit Chupeng, 
Bukit Lagi, Bukit Wang Tangga and Tebing Tinggi (Perlis). More than 700 numbers, 


Holttum, R.E. Oct. 3rd 1922; Gua Batu, Selangor, several ferns. Aug. 22nd-26th 1925; 
Langkawi, around Pulau Dayang Bunting and Pulau Timun. About 25 numbers. 

Johnson, A. Nov. 17th 1968; several ferns from Gua Batu, Selangor. 

Kadim, Tassim. Aug. 13th 1959 (with Allen, B.E.G.M.); several numbers from Gunong 
Tempurong, Perak. 

Kassim, Rajab. Aug. 17th 1962; several numbers from Gua Batu, Selangor. 

Kelsall, H. Jan. 1891; several plants from Gua Batu, Selangor; all poorly labelled. 

Keng, H. Sept. 25th 1954; at Kuala Trengganu, several ferns. Mar. 28th 1962 (with 
Mrs. Keng); a specimen of Monophyllaea horsfieldii R. Br. Nov. 3rd—7th. 1968; Lang- 
kawi islands (sometimes with others); some assorted specimens. 

Kerr, A-F.G. June 20th 1932; Langkawi, around Kuah; several numbers. 

Kiah, Hj.M.S. April 11th-May 9th 1938: collected on Gunong Baling, Kedah and Kaki 
Bukit, Perlis. More than 150 numbers. 

King, Sir G. See under Kunstler. . 

Kunstler, H.H. (collector for Sir G. King, as King’s collector) 1880-1885; collected around 
Gopeng and Larut, also visited Gunong Pondok (all in Perak). Almost 100 numbers. 


Loh, H.S. July 1970; Gua Peningat, Pahang. Recorded 101 numbers. This is the first known 
occasion that this outcrop (at 713 m the tallest in Malaya) has been climbed. February 
1972; visited Gua Musang, Kelantan. Sixteen numbers. 


Mahmud, S. 1970-1971; Bukit Anak Takun (Selangor), and Ipoh (Perak). Several numbers 
See also Samsuri. 


Mat ona W. Jan. 1939; a specimen of Cymbogon calcicola Hubb. from Gunong Baling, 
edah. 


Merton, L.F.H. Dec. 22nd 1960; Gua Batu, Selangor; several numbers. 

Mills, G.R. April 16th 1925 (with Henderson); Gunong Lanno, Perak; several specimens. 

Nauen, J.C. Nov. 13th-17th 1941; Gunong Baling and Langkawi, both in Kedah. About 
20 numbers. See also Corner. 


Ng, F.S.P. Oct. 1966-Feb. 1968; Gunong Mesah, Gunong Rapat, Gunong Tempurong and 
other hills around Ipoh (all in Perak). Gua Batu and Bukit Takun in Selangor and Gua 
Musang in Kelantan. More than 150 numbers. 


Nur, Mohamed. 1931-1937; Bukit Sagu, Pahang and Bukit Takun and Gua Batu, Selangor. 
See also Haniff. About 160 numbers. 


170 Gardens’ Bulletin, Singapore — XXX (1977) 


Ogata, K. Feb. 14th-15th, 1968; Gunong Gajah and Gunong Tempurong in Perak. Some 
assorted specimens. 
Phang, C.I. Sept. 15th 1960; several ferns from Bukit Anak Takun, Selangor. 


Poore, M.E.D. Sept. 12th 1960 & July 16th 1961; visited Gua Batu, Selangor. Several 


numbers. 
Reid, J. Jan. 29th 1950; three numbers from Bukit Takun, Selangor. 


Ridley, H.N. 1896-1897; visited Kuala Dipang (Perak) and Gua Batu, Selangor 1898-1920; 
visited Gua Batu about five times, also Gunong Keriang, Kedah and Bukit Lagi, Perlis. 
About 150 numbers. 


Samat, A. 1961-1969; visited Langkawi (Kedah), Bukit Anak Takun and Bukit Takun 
(Selangor). Mainly ferns; about 15 numbers. 


Samsuri, A. Mar. 8th-14th, 1971 (with Mahmud); collected in Perak, around Gopeng, on 
Gua Putri, Gunong Pondok, Gunong Rapat, around Ipoh, and in Tambun district. About 
120 numbers. 


Shimizu, T. Oct-Nov. 1967 (with Stone); visited Gua Musang (Kelantan) and Bukit Takun 
(Selangor). Some ferns. 


Sinclair, J. 1958-1962; visited hills around Ipoh (Perak) and Gua Batu (Selangor). About 
30 numbers. 


Singh, Hardial. Jan. 23rd 1966; visited Gua Batu (Selangor). Several numbers. 
Smith, G. One Chirita caliginosa Cl. from Perak Cave Temple. 

Smith, J.W. July 19th 1956; one Vitex siamica Will. from Gunong Baling, Kedah. 
Soepadmo, E. Sept. 1968; one Carallia brachiata (Lour.) Merr. from Ipoh, Perak. 
Spare, G.H. Oct. Ist 1939; Gunong Pondok, Perak. Several numbers. 


Start, A. Jan. 23rd 1972; one Mucuna biplicata Teysm. et Binn. from Bukit Anak Takun, 
Selangor. 


Stone, B.C. 1965-1972; visited Langkawi (many of the islands), Kedah, Gua Musang 
(Kelantan), Gunong Pulai (Perak), Bukit Anak Takun, Bukit Takun and Gua Batu 
(Selangor). Chintamani (Pahang). About 450 numbers. 

Students (University of Malaya 1967) May 1967; some numbers from Langkawi (Kedah). 

Symington, C.F. 1934-1938; visited Langkawi (Kedah) and Bukit Takun. About a dozen 
numbers. 

Teruya, Z. Jan. 20th 1929; one Monophyllaea horsfieldii R. Br. from Gua Batu, Selangor. 

Tomlinson, P.B. Sept. 1955; one Monophyllaea hirticalyx pr. from Ipoh Perak. 

Turnau, E.A. Aug. Sth 1962; two numbers from Langkawi, Kedah. 

UNESCO 1962 July 27th-Aug. 13th, 1962; visited the Kelantan limestone around Gua 
Musang and Bertam, collected on Batu Bayan, Batu Hayan, Batu Ner, Batu Pinta, Gua 


Batu Boh, Gua Musang, Gua Serai, Gua Seri and Gua Panjang. About 700 numbers. 
Led by Dr. H. Keng. 


Whitmore, T.C. 1966-1970; visited Langkawi (Kedah), Gua Batu Boh, Gua Musang, Kuala 
Jenera, Sungei Nenggiri (Kelantan), and Bukit Anak Takun, Bukit Takun, and Gua 
Batu (Selangor). About 120 numbers. 


Wong, Y.K. May 13th 1962; a Maxburretia rupicola (Ridl.) Furt. from Bukit Takun, 
Selangor. 


Wooldridge, T.A. June 1896; a Dracaena curtisii Ridl. from Langkawi (Kedah). 

Wray, L. 1894; two specimens from Perak limestone; labels mutilated, one undated. 
Wyatt-Smith, J. June 11th 1960; one Salacia grandiflora Kurz. from Gua Batu, Selangor. 
Wycherley, P.R. Dec. 28th 1969 (with Stone); Gua Batu, Selangor. Some assorted specimens. 


__ For the purpose of this study most of the limestone areas in Malaya were 
Mim and numerous hills explored and botanised. About 1550 numbers were 
collected. 


The plants treated include the terrestrial as well as the epiphytic members. » 
Included also are species from hill bases where the soil is definitely of limestone 
Origin; this is usually a rusty-red loam and very distinctive. Characteristically there 
are numerous boulders strewn about. Admittedly it is often difficult to decide what 
does or does not constitute the limestone vegetation. Certainly the vegetation at 
hill bases (with limestone derived soil) is different from that on hill tops, but 
equally it also differs from that of the surrounding lowland forest. This hill base 


Malayan Limestone Flora 171 


vegetation has arisen as a result of the interaction between the presence of the 
hill and the surrounding forest. Accordingly I have included the species found 
here in this treatment. 

In the case of plants from the small outcrops recently discovered in Johore 
(Rajah, 1970) great care was exercised to see that only those actually growing on 
the rocks were recorded. This is because the outcrops here are very limited and 
low, from boulder size protrusions to several of about 20 m tall and as broad. 
Most of the cracks and crevices are filled with soil derived from the surrounding 
sandstone formation. (This publication also records the first botanical exploration 
on the Johore limestone.) 

All specimens from ‘near the base of hill’, or from ‘hill bases’ as recorded 
on the labels of herbarium sheets are excluded unless the labels also specify that 
the particular plant was from limestone rock or soil. 


In order to obtain as complete a record as possible of the limestone flora, 
every sheet in the herbaria at the Singapore Botanic Gardens, the University of 
Singapore and the University of Malaya was scrutinized. A start was also made 
at the herbarium of the Forest Research Institute but as the search was not 
showing success it was stopped. Practically all the old and most of the new lime- 
stone records there (at Kepong) are duplicated in the Singapore Botanic Gardens 
herbarium. Subsequently the examination of specimens at Kepong was restricted 
to the known limestone genera and to recent (post 1960) records from limestone 
districts (with the help of collector’s field notes). 

Many labels are unclear and to decide whether a specimen was recorded from 
limestone or not is not always straightforward. Sometimes the town, village or 
district bears the same name as the limestone hill (or vice versa), e.g. Baling, 
Gua Batu and Gua Musang. Specimens in such cases were excluded unless the 
record states explicitly that it was from the hill. 

By searching the herbaria and literature I have managed, I think, to include 
all the species (possibly with few exceptions) so far recorded from the Malayan 
limestone. 


Purpose and Scope 

This work attempts to present a comprehensive account of all pteridophytes 
(true ferns only) and spermatophytes ever found growing on limestone hills and 
sites in the Malayan Peninsula. Included also are the non-indigenous species 
(introduced, inadvertently or otherwise) which have become naturalized and now 
form part of the wild-growing population. 

Sect. 1 discusses the geology, origin and distribution of the limestone hills, the 
vegetation, phytogeography and size of the flora, and affinities of the floristic 
components to limestone. 

Sect. 2 the flora proper, starts with the Pteridophytes (true ferns only), going 
on to the Gymnosperms and ending with the Angiosperms (Dictoyledons and 
Monocotyledons). Under these four main groups, the families are arranged in 
alphabetical order and thereunder the genera and species. Dichotomous keys 
leading to the identification of almost all the species are included. For the 
exceptionally large groups, the Pteridophytes, Euphorbiaceae, Orchidaceae and 
Rubiaceae, an introductory key with leads to the main key is provided. 


172 Gardens’ Bulletin, Singapore — XXX (1977) 


Every species is provided with its approved name and the original place of 
publication. Synonyms, if any, especially those published in major works relevant 
to the local flora, are cited. References are also made to publications on the local 
flora, and reference to Henderson (1939) is cited for every species included in 
his work. 


The names of characteristic species (those that come under my ‘affinity’ 
grouping I and IJ in ch. 5) are printed in bold face type. In addition, brief notes 
on the distribution, and frequency on the Malayan limestone are often included for 
these species. 


All literature cited in the discussion are listed in full in the bibliography at 
the end of this work. 


Specimens are cited only in special cases, such as rarity, or doubtful locality, 
or in relation to certain taxonomic problems. However, full lists of specimens are 
to be found in the dissertation paper, and this may be consulted in the Library 
of the University of Malaya or in the University Herbarium, Kuala Lumpur. 


Lastly Bryophyte collections were made during this study but are not included 
here because of the difficulty in obtaining identifications. A further study of this 
group would be desirable. 


2. GEOLOGY, ORIGIN AND DISTRIBUTION OF THE LIMESTONE 
HILLS. 


Geology 


Most of the limestone hills in Malaya rise from flat or undulating plains. They 
form conspicuous and often spectacular hills up to 2342 ft. (713 m.), (Gua 
Peningat, Pahang), often with sheer, vertical or overhanging cliffs. 


They range in age from Ordovician to Triassic. The estimation of age has 
been a difficult problem since are recrystallised, unfossiliferous (Hutchison, 1968), 
and as all studied samples are recrystallised, thermoluminescence cannot be used 
to differentiate limestone stratigraphically in Malaya (Hutchison, 1968). Up to 
as late as the 1950’s anything calcareous in Malaya has been assumed to be 
carboniferous and was ascribed to the calcareous formation. Since then many 
studies have been made, providing a more accurate picture of the age of our 
limestone, e.g. Paton (1961), Ingham and Bradford (1960), Gobbett (1965a), 
Hutchison (1968) and Suntharalingam (1968), but the ages of many still remain 
indicated by only a very rough estimation. 


The Kedah/Perlis(including Langkawi) limestone areas are mainly Ordovi- 
cian-Silurian. The Selangor limestone appears to be confined to the Silurian 
(Gobbett, 1964), and the recently discovered Johore limestone is Permian (Rajah, 
1970). Most of the others are Permian (Paton, 1961), but further studies will 
probably show a variation in age from the Silurian to Permian (as is the Selangor 
limestone Paton included under Permian). However no hill has been found to be 
Triassic which so far is limited to small lenticular beds in Pahang (Paton, 1961). 


It is generally thought that the older limestone gives rise to more or less 
continuous ranges of hills without marked vertical cliffs whereas on the other hand 
the younger limestone produces more isolated hills with high, vertical or even 


overhanging cliffs (Paton, 1961). This weathered-look is shown by the Kedah/- 


Perlis (including Langkawi) limestone of Ordovician-Silurian age. This genera- 
lisation was. made before the Selangor limestone was known to be Silurian, 
and now no longer holds. This is because the latter shows marked cliff development 
(except for Bukit Anak Takun). Thus it would seem that the age of the actual 
limestone rock itself has little bearing on the development of the hills, and there is 
no difference in form between hills from Permian and Ordovician-Silurian rock. 


Malayan Limestone Flora 173 


Our limestones are generally very pure. Most are white, pale grey or slightly 
yellowish; some are dark grey to almost black because of carbonaceous or 
argillaceous impurity. Rarely it is red because of hematite inclusion or by iron 
staining. Hutchison (1968), who analysed 182 very random samples chemically 
(which therefore should give a rough indication of the true nature of the Malayan 
limestone) recorded that most of them have about 2.5% insoluble residue with an 
arithmetic mean of about 4.4% insoluble residue. He further notes that 81% of 
his samples are limstone (CaCO,) and 19% dolomite (CaMg (COs) :). 


Origin 

Originally sedimentary, the Malayan limestone is, as mentioned earlier, 
recrystallised. It is thus in fact marble. This happened when peninsular Malaya 
underwent an orogeny (an episode of mountain-building during which the rocks, 
including the Malaysian limestone were compressed, folded and heated) in the 
Triassic period (about 200 million years ago). Towards the end of the Triassic 
orogeny the rocks were uplifted into hills and mountains. The botanical stage was 
set, since then most of the Malayan limestone was never submerged again, and 
they were ready to receive their first plant. However it is not certain how long it 
took before the limestone topography as we know came into being. 


Various theories have been put forward to explain the existence of the lime- 
stone in the form of spectacular karst-tower hills. These are reviewed by Paton 
(1964), and include block faulting, sub-aerial erosion, folding, marine erosion and 
plastic flow. The most plausible explanation seems to be simple sub-aerial denuda- 
tion modified in certain cases by marine erosion. 


The theory of sub-aerial erosion takes into account the high solubility of 
calcium carbonate compared to the majority of the silicates, of which most of all 
other rocks are composed. This is an important point, for any theory seeking to 
explain why the limestone hills are in the form they are must also explain why 
formations of other rocks are not existing in the form the limestone hills are. 
Weathering of silicate rocks produces more or less rounded topography; on lime- 
stone deep channels are often cut. 


Streams and swamps at the bases of limestone (assuming that they were once 
well-jointed and continuous) are highly effective in eroding it. The pH of swamp 
and stream water in low-lying areas is commonly as low as 3.5 (Paton, 1964). 
This acidic water gradually erodes the rock, forming channels; floors and walls 
are enlarged, and from above, downward erosion cuts deep channels that 
eventually reach the base. The underground channels continue to expand laterally, 
vote resulting in the collapse of the roofs, producing steep-sided and isolated 

S. 


The initial stages of erosion by water are evident in the form of grooves or 
nips, characteristic of many coastal limestone areas in Langkawi and at the bases 
of hills on the mainland. Some grooves on hills in the Kinta and Kedah/ Perlis 
area are marine in origin, formed in the post-glacial periods (post Pleistocene) when 
the sea level, at least 50 ft. higher than now (Scrivenor, 1949; Walker, 1956), 
flooded the plains. 


Distribution 

Until recently, all the known limestone outcrops were from north of Kuala 
Lumpur. However in 1965-66, officials of the Geological Survey of Malaysia 
discovered limestone outcrops in Johore while carrying out geological mapping of 
that area (Rajah, 1970). The outcrops are in an area 500 by 1200 m on the south- 
eastern slope of Gunong Sumalayang at an elevation of 250 to 400 m. There are 
about 30 outcrops here varying in size from boulders 1-2 m across to the largest 
about 20 m tall and as wide. 


174 Gardens’ Bulletin, Singapore — XXX (1977) 


Elsewhere in Malaya the limestone hills are in their most spectacular and 
prolific development in south-west Kelantan and north-west Pahang, in the Kinta 
valley from Kampar northwards to Ipoh, Tambun and Tanjong Rambutan, and 
in north-west Perlis. 


Hills are also found around Alor Star, Kedah, elsewhere in Pahang, near 
Kuala Lumpur, Selangor and rarely in Trengganu. They are very prominent in 
the Langkawi group of islands off the extreme north-west coast of Malaya. Not 
all the islands are limestone, and some are limestone only in part, but generally 
limestone is abundant, especially in the eastern and southern aspects of Langkawi. 
This is the only coastal limestone in Malaya. 


The distribution of the Malayan limestone hills is shown in Maps 1, 2 & 3. 
All the hills botanised are included in the maps except some of the smaller hills 
which are not named even in regional 1 inch to 1 mile maps. It has also not been 
possible to include all existing outcrops, but all the limestone localities are 
represented and most of the popuarly and less popularly known have been included. 


3. VEGETATION 


The limestone vegetation forms a characteristic and easily recognised forest 
type in Malaya. It is a distinct Edaphic Climax Formation (Symington, 1943). In 
fact, Corner (1960) considered the limestone forest as one of the three constituent 
forest types (the other two being the lowland and the mountain forest). 


The vegetation supports an extremely diverse and rich flora which is a 
reflection of the diversity of the habitat. Henderson (1939) classified the Malayan 
limestone hills into three broad groups, viz.: 

1. Very dry hills with little soil and much exposed rock. Trees usually 
rather stunted; mosses and herbaceous plants scarce. 

2. Wet hills often with gullies filled with rich soil. Trees fair-sized and not 
stunted; mosses and herbaceous plants present. 

3. Soil-rich hills, well covered with soil (usually a stiff red clay), little rock 
exposed. Trees tall; mosses and herbaceous plants few. 


This is in effect, as Henderson further discussed, dividing the limestone hills 
into two general categories, ‘dry hills’ (1) and ‘wet hills’ (2 and 3). This classifi- 
cation however does not do justice in summing up the diverse limestone habitat. 
‘Dry hills’ often have sheltered gullies with soil supporting fair-sized trees, mosses 
and herbaceous plants. Similarly ‘wet hills’ often have exposed rocky ridges with 
stunted vegetation. 


Anderson (1965) subdivides the Sarawak limestone habitat into eight groups, 
viz.: 1. Alluvial soils at the base of hills. 2. Base of hills and ravines between hills. 
3. Slopes of hills 4. Cliffs. 5. Scree slopes on hills. 6. Summits of hills at low 
altitudes. 7. Submontane limestone. 8. Secondary vegetation on hills. 


In Malaya there are no submontane limestone hills and this classification 
does not seem to effectively describe the Malayan limestone habitat. Thus 
another classification is proposed. This is, like the other schemes, only provisional 
and there is a great deal of overlap between the subdivisions. This subdivision 
of the habitat takes into account the geology and topography of the hill, the 
physiognomy of the vegetation and the floristic composition. Succession is also 
considered and disturbed or secondary vegetation is therefore made a subdivision. 


Malayan Limestone Flora 175 


PULAU 
TIOMAN 


cea Ss ( 
= 


JOHORE 


GUNONG® 
SUMALAYANG 


MAP 1. MALAYA IN GENERAL 


176 


6° 20S 


Gardens’ Bulletin, Singapore — XXX (1977) 


199° 50°E 


PULAU LANGKAWI 


199° 50'E 


MAP 2. LANGKAWI, SHOWING THE MAJOR LIMESTONE LOCALITIES 


Key to the numbers (in a clockwise direction starting from the north). 


1. Pulau Dangli 11. 
2. Pulau Gasing 
3. Pulau Pasir 12. 
4. Tanjong Batu Kulat 13. 
5. Tanjong Ru 14. 
6. Gua Cherita 15. 
7. Pulau Langgun 16. 
(with lake) 17. 
8. Pulau Tanjong Dendang 18. 
9. Tanjong Dagu 19. 
10. Tanjong Dagu Forest 20. 
Reserve 21. 


Selat Panchor Forest 
Reserve 


Selat Panchor (straits) 
Pulau Timun 

Pulau Lading 

Pulau Chupak 

Pulau Bumbon Besar 
Pulau Bumbon Kechil 
Telok Sambar 
Tanjong Tirai 

Pulau Dayang Bunting 
Pulau Jerkom Kechil 


22. 


Pulau Jerkom Besar 
Pulau Singa Kechil 
Pulau Singa Besar 


Tasek Dayang Bunting 
(lake) 


Pulau Beras Basah 
Pulau Rebak Besar 
Tanjong Belua 
Tanjong Chinchin 
Tanjong Tok Manap 


Ly 


Malayan Limestone Flora 177 


The subdivisions are: 


Base of hills 

Talus slopes 

Hill slopes to about 60° steepness 

Gullies and valleys 

Cliffs and near-vertical slopes 

Summits with considerable soil cover 
Summits with none or very little soil cover 
Coastal limestone 

Disturbed areas 


CHONAARWN 


Although the nature of the habitat necessitates a classification taking into 
account all the characteristics present to separate the subdivisions effectively, this 
classification is more physiognomic than floristic in kind. The physiognomy of the 
vegetation is in turn dependent on the topography of the hill, the abundance or 
scarcity of soil, moisture and shelter. The floristic composition however varies a 
great deal from hill to hill and although it would be possible to subdivide a hill 
floristically, this pattern of floristic distribution would not hold for other hills. 
Species characterising a dry rocky summit like Bukit Takun (Buxus malayana and 
Wikstroemia androsaemifolia) are not found on a similarly dry and rocky part 
of, for example, Batu Neng. Sycopsis dunnii is common on Bukit Serdam summit 
(with thin soil) but on Gunong Rapat summit (also with thin soil) a common 
plant is Murraya paniculata, and Sycopsis dunnii is absent, and the converse 
applies. 

Floristically also, the limestone north of Alor Star, including those in 
Langkawi, mainland Kedah and Perlis differs from that further south. They form 
part of the Thai flora which begins north of Alor Star, in Kedah (Ridley, 1911) 
or north of the mouth of Kedah river which is near Alor Star (Kloss, 1922). (This 
difference in the floristic composition is due mainly to the regular dry season from 
December to February which is confined to the extreme north of Malaya). 


However in this classification, the Langkawi limestone is treated in the same 
manner as the rest and not as a distinct floristic entity. This is in order to provide 
a coherent picture of the Malayan limestone in which floristic composition is one 
of the criteria used and not the deciding factor. 


The examples of plants cited in the following discussion are representative of 
the subdivisions of limestone habitats in which they are found. However a con- 
siderable number of such examples do not have rigid habitat preferences and may 
also be found in the other subdivisions. Though they are representative of the 
various subdivisions they are not necessarily found on all hills with similar 
habitats; in fact many species form very local populations on one or several hills. 


1. Base of hills 


The bases of hills are usually covered by a mixture of alluvial soil not derived 
from the limestone but from the surrounding geological formation and also soil 
derived from weathering of the hills. The vegetation here varies with the locality 
of the hill and probably with the nature of the derived alluvial soil (from the 
surrounding geological formation). Protected valleys (wangs) may occur; these 
are surrounded, often entirely, by abrupt limestone walls. The floor is usually the 
typical Terra Rossa soil. Such areas are often cultivated (as in the hills around 
Ipoh). 

The vegetation is generally tall and closed, with the crowns of the plants 
touching or overlapping. Typical species include Villebrunea sylvatica, Gmelina 
asiatica, G. villosa, Dendrocnide stimulans, Diospyros cauliflora, Mallotus 
philippensis, Melanolepis multiglandulosa, Orophea hirsuta and Pisonia umbelli- 
fera. Some climbers are also found e.g.; Jodes cirrhosa and Mucuna biplicata. 


178 Gardens’ Bulletin, Singapore — XXX (1977) 


Sometimes where water drips down from steep rocky slopes or from over- 
hanging cliffs, a distinct herb community develops. Species include Alocasia lowii, 
Chirita caliginosa (which is also found on dry situations) Epithema saxatile and 
Monophyllaea horsfieldii. Bryophytes are usually luxuriant. 


2. Talus slopes 


At and near the base of many hills are accumulations of boulders, organic 
matter and general debris from the hill. Rocks from this scree have evidently 
fallen from the hill. Often the bases of hills surrounding amphitheatres or wangs 
have talus slopes at their bases. These are probably remnants of the roofs of large 
caverns that have suffered a cave-in. The vegetation here is tall and closed, 
physiognomically not unlike that on cliff bases (Gp. 1.). The composition is 
however somewhat different; and perhaps the distinguishing character is the herb 
and shrub vegetation characteristing such a shady, rocky terrain. The trees and 
shrubs include Atalantia roxburghiana (peculiar to Bukit Takun), Cleidion 
javanicum, Morinda elliptica, Streblus ilicifolius and Trivalvaria macrophylla. The 
herbs include Heterogonium pinnatum, H., alderwereltii, Impatiens spp. (1. mirabilis 
dominates the scenery on Pulau Langgun, Langkawi, north of the lake), Pilea 
and Procris. 


3. Hill slopes to about 60° steep 


These more gentle slopes are nearly always covered with soil and support a 
dense, close vegetation 5-10 m or more tall. The soil may be thin or to about 
1 m deep at places, and boulders are usually present emerging from the soil. 
Ground vegetation is usually spares and is often associated with the boulder out- 
crops. These include Antrophyum parvulum (on rocks), Ophiorrhiza, Piper, and 
in the more sheltered niches Gymnostachyum decurrens and Scutellaria discolor. 
The trees and shrubs include Cladogynus orientalis, Croton cascarilloides, Mallotus 
dispar, Microdesmis casearifolia, Orophea cuneiformis, Polyalthia brunneifolia, 
Saraca declinata, S. thaipingensis and Rinorea bengalensis. On Gua Kechil, Raub, 
the rare tree Diospyros adenophora is fairly common. 


4. Gullies and valleys 


These are often the most sheltered spots on hills. This group includes 
pot-holes, depressions, gullies, valleys, ravines and other concavities which may 
be narrow and small or broad and extensive, extending between two peaks on the 
same hill. The ground is usually deep or shallow soil with a covering of litter and 
other organic debris. The vegetation is fairly dense, 5-10 m tall with occasional 
emergents, and physiognomically not unlike that on soil-covered slopes and talus 
slopes. The floristic composition varies somewhat and its location and topography 
is usually distinctive. (Sometimes however gullies and valleys may emerge into 
hill slopes.) Gullies may even be present on the so-called ‘dry hills’ (the most 
outstanding example is Bukit Takun), harbouring mosses and delicate herbaceous 
plants like Corybas mucronatus and Goodyera hispida. The ground vegetation is 
usually quite sparse, no doubt because of the closed canopy. Bryophytes are 
common and herbs like Epithema saxatile and Monophyllaea are usually associated — 
with boulder outcrops. 


The trees and shrubs include A grostistachys gaudichaudii, Canthium didymum, 
Fagraea curtisti, Randia densiflora, Sauropus suberosus and Sterculia rubiginosa. 
Climbers like Connarus and Lasiobema are found and the palms present include 
Aaa westerhoutii (also from talus slopes), A. hookeriana and Iguanura poly- 
mor pha. 


Malayan Limestone Flora 179 


_ EN 


MILES 


———— International boundary 
———— Stote boundary 
seer Railway 


ot ere cies aby 


“i . 
vl 


Bertam 


e 
Batu Pintg 


- Kuala @ 8 
J Bati 

Gunong ‘s NGua Panjang 
Pondck 


{ ae i 
7 ao Pes 
be bel RAMBUTAN -—— ; 
4 eT, N>\ v j rere ue Layers Ve” 
.Gue Peningat 


coh 
yap 
exRnt? x 


-KUALA 
LUMPUR 


100° 101° 102° 103° 


MAP 3. MALAYA (EXCLUDING LANGKAWI!) SHOWING THE MAJOR LIMESTONE OUTCROPS 


Key to the numbers indicating position of outcrops. 


1. Bukit Besih Hangat 5. IPOH — ‘including Perak Cave Temple 
Bukit Wang Tangga 6. GOPENG — including Gunong Lanno 
Tebing Tinggi Gunong Rapat 

2. KODIANG — including Batu Hantu 7. Kuala Dipang 

Batu Kalong Sungei Siput 
3. LENGGONG — including Batu Tukang g GUA MUSANG — including Batu Machang 
Gua Badak Batu Papan 
Gua Pipit Batu Tapah 
Gua Putri Gua Serai 


4. TAMBUN — including Gunong Ginting 
Rotan Segar 


180 Gardens’ Bulletin, Singapore — XXX (1977) 


5. Cliffs and near-vertical slopes 


Vertical cliffs, overhanging cliffs and near vertical slopes present a unique 
habitat and support a distinct vegetation which depends on the degree of exposure 
of the cliffs, presence or absence of soil pockets and availability of moisture. This 
is a very easily recognised subdivision topographically and the vegetation varies 
from very sparse and open (or sometimes absent) to a fairly dense herb and scrub 
cover. This denser vegetation cover is found along ledges where there is accumu- 
lated soil and debris and in other soil pockets. Where there is seepage or water 
dripping down a characteristic vegetation develops along this line of moisture. 
This may be on overhangs or clefts on cliff faces. If on clefts this vegetation 
decends to the base of the cliff and is very similar to that described under sub- 
division 1 (where water is available). This includes Chirita, Epithema saxatile, 
Monophyllaea and the fern Adiantum malesianum (which is also from dry 
localities) and bryophytes. 


Cliff faces with no cracks or crevices do not support any macro-vegetation, 
but those with cracks and crevices which invariably collect soil and other debris 
support a distinctive flora. The species include the very characteristic Boea, 
Paraboea and Chirita. Boea spp. especially are often very prominent high up on 
cliff faces. 


Near vertical slopes and cliffs with ledges and other soil pockets often support 
a herbaceous or scrubby vegetation. The species include Canscora pentanthera, 
Cymbopogon calcicola, Callicarpa angustifolia, Cheilanthes farinosa (only on 
Bukit Chintamani), Jasminum, Pyrrosia stigmosa and Zizyphus oenoplia. 


6. Summits with considerable soil cover. 


Most hill summits are covered with soil (at least in part). Some have 
considerable to extensive rock outcrops with sparse vegetation (these rocky spots 
are included under subdivision 7.) In places scattered boulders and pot-holes are 
common; generally the topography is uneven to very rugged. Ridges with some 
soil supporting a closed vegetation are included here. The vegetation is usually 
scrubby, 2-7 m tall and the canopy is closed. In places the plants are spindly. 
Generally these shrubs and trees are fairly well spaced, and though the canopy 
is closed there usually is a rich ground flora. Occasional emergent trees are present, 
eg; Mangifera sp. on Gunong Pondok, Madhuca ridleyi on Bukit Serdam and 
Garcinia murdochii on Gua Kechil. 


The trees (they are usually somewhat stunted, though not extremely so) and 
shrubs include Cleistanthus gracilis, Decaspermum fruticosum, Dehaasia curtisii, 
Eriobotrya bengalensis, Erythroxylum cuneatum, Eugenia pendens, Garcinia 
minutiflora, Mallotus brevipetiolatus, Mallotus miquelianus, Murraya paniculata, 
Neolitsea zeylanica and Sycopsis dunnii (especially on Bukit Serdam). The pandan, 
Pandanus irregularis is common and conspicuous on ridges on Batu Neng. 
Climbers and scramblers are usually scarce and include Maytenus curtisii and 
Lygodium polystachyum. 

The ground vegetation is fairly rich in bryophytes especially in the more 
sheltered spots on rocks and at the basal parts of tree trunks. The herbs include 
the lycopod, Lycopodium squarrosum; ferns are common and in places from the 
dominant ground cover; they include Asplenium adiantoides, A. macrophyllum, 
Loxogramme avenia, Photinopteris speciosa, Phymatodes scolopendria and 
Pyrresia stigmosa. Ground orchids are equally common and include Agrostophyl- 
lum bicuspidatum (which may also be a low epiphyte), Coelogyne asperata 
(which on parts of Batu Neng and Batu Tapah forms almost pure mats), 
Dendrobium luxurians, Eria rigida and Renanthera histrionica. Epiphytes are 
usually not conspicuous though occasionally they are found; these include orchids, 
e.g. Appendicula torta and Bulbophyllum apodum and ferns, e.g. Pwrosia. 


Malayan Limestone Flora 181 


7. Summits with no or very little soil cover 


Most hills have some exposed rocky parts, while a very few have extensive 
rocky summits. Soil is scanty and found in little hollows, cracks and clefts in the 
rock but is never completely absent. Even extensively rocky summits have gullies 
and valleys where soil and litter are accumulated and the vegetation supported is 
different (included under subdivision 4.) 


The vegetation on these rocky summits is sparse and open but fair-sized 
trees rooted in clefts and hollows are present. Most are, however, stunted and 
gnarled though occasionally large ones occur. An example are the figs, the roots 
of which ramify all over the rocks, into every crack and crevice and often 
extending for many meters over cliff faces in an intermeshed network. 


Bukit Takun in Selangor is the classic example of a hill with an extensive 
rocky summit, but other examples occur, e.g. in Langkawi. The plants on these 
hills include some herbs usually from soil pockets and from amongst the surface 
roots of trees where some debris have accumulated. These include Adenia nico- 
barica, Amorphophallus spp., Arisaema fimbriatum, A. roxburghii, Boea spp., 
Chirita spp. and Dichanthium mucronulatum. It is interesting to note that of these, 
Adenia nicobarica, Amorphophallus spp. and Arisaema spp. have fleshy tubers 
which often grow into cracks and crevices. The possession of tubers could be an 
advantage under such dry and exposed conditions. Climbers present include 
Dischidia hirsuta, Morinda umbellata and Secamone micrantha, and the pandan, 
Pandanus alticola (commonly an epiphyte outside the limestone field) which is 
often present growing from rocks (it is abundant on Bukit Takun). 


The trees and shrubs present include Buxus malayana, Diospyros ferrea 
(commonly on hills around Gua Musang), Ficus calcicola, Glycosmis calcicola, 
Memecylon laevigatum, Pistacia malayana, Planchonella obovata, Podocarpus 
polystachyus, Tarenna curtisii, Terminalia triptera (only Langkawi), Vaccinium 
littoreum (only on Bukit Takun and Anak Takun), and Vitex siamica. 


8. Coastal limestone 


In Malaya this only occurs in Langkawi where part of the coast of the main 
island and many of the smaller islands are entirely limestone. Hills and islets 
which descend gradually to the sea are often fringed on the seaward edge by 
narrow sandy beaches (non-limestone, and derived from other geological forma- 
tions as weathered limestone produce hardly any ‘sand’) on which develops a 
typical strand vegetation. It is these low limestone areas that are mostly influenced 
by the sea and are invaded by some species from the strand vegetation. Hills with 
tall abrupt cliffs do not show any littoral influence. However, it must be remember- 
ed that many species from the Langkawi limestone are not found on limestone 
further south on the mainland, but this is due to the northerly position of 
Langkawi principally, and not due to littoral influence (which is coincidental and 
at the most contributary). These species include Bombax anceps, Cycas siamensis, 
Colona merguensis, Euphorbia antiquorum, Grewia viminea, Hopea ferrea, 
Pentacme siamensis, Sterculia lancaviensis and Vatica cinerea. 


The coastal limestone therefore is restricted to that part of the limestone in 
close proximity to the sea, and this is usually near sea-level. The seaward edge is 
fringed with sand which supports a typical Indo-Malaysian littoral vegetation 
(sensu Schimper, 1903). The vegetation on this limestone is sparse to dense, short 
or tall, depending on the availability of soil and the topography; but littoral 
species are present. The species include some herbaceous members like Davallia 
solida, Humata pectinata, Eurycles sylvestris and Tacca leontopetaloides. Trees 
and shrubs include Barringtonia asiatica, Caesalpinia crista, Guettarda speciosa, 


182 Gardens’ Bulletin, Singapore — XXX (1977) 


Heritiera littoralis, Pandanus odoratissimus, Peltophorum pterocarpum, Thespesia 
populnea and Xylocarpus granatum. Together with the species mentioned above 
this makes up a distinctive coastal limestone vegetation (probably distinctive even 
if there were other coastal limestone areas further south). 


9. Disturbed areas 


Disturbed areas usually occur on the summits, lower slopes and bases of hills. 
These disturbances could have been brought about by a number of causes, viz: 


(i) Fire, occurring naturally or artificially, e.g. Bukit Takun in August 1970 
and Gua Musang in mid-1969. 


(ii) Mining and quarrying activities. This is seen on a large number of hills 
in all the major limestone districts. 


(iii) Agriculture, affecting only hill bases, e.g. some hills around Ipoh and 
in limited spots on many other hills. 

(iv) Erection of communication transmitters and trignometrical stations, e.g. 
on Gunong Lanno, G. Layang-Layang and G. Tasek, also on other hills. 


(v) Frequent trampling by visitors, e.g. on Bukit Takun and Gua Musang. 


Whatever the cause, disturbances result in the partial or complete destruction 
of the original vegetation. It is interesting to note that Cratoxylum maingayi, 
Podocarpus polystachyus and Radermachera lobbii survived the 1969 fire on 
Gua Musang that destroyed all the other vegetation affected. Cratoxylum and 
Podocarpus probably survived because they were the largest trees there and con- 
sequently had thick bark. Several plants of Radermachera (they were slender and 
probably not more than 4 m tall) were burnt almost to the ground. By August 
1971 the sucker-shoots sent out were already 2 m tall and flowering. 


Though the complete development from sere to high forest has never been 
observed, a composite and representative picture (at least of the initial stages) 
can be formed by observation of a number of hills. One of the earliest colonisers 
is the moss Bryum coronatum Schwaegr. which was abundant on Gua Musang 
in August 1970 and also in August 1971 on burnt earth and on rocks. This moss 
is followed (or accompanied) by a large number of ubiquitous weeds, truly 
opportunistic and very adaptable and probably completely indifferent to the lime- 
stone influence. The species include ChArysopogon aciculatus, Colocasia gigantea 
(prominent on Gua Musang), Fleusine indica, Erechtites valerianifolia, Eupato- 
rium odoratum, Macaranga tanarius, Muntingia calabura (prominent on _ the 
quarried slopes of Bukit Chintamani), Nephrolepis biserrata, Neyraudia reynau- 
diana (conspicuously gracing the summit of Gua Musang in 1971), Paspalum 
conjugatum, Pityrogramma calomelanos, Pteridium caudatum var. yarrabense, 
Pteris vittata, Rhynchelytrum repens and Solanum ferox. Limestone elements 
present at this early stage include Boea spp., Chirita spp. and Schizaea inopinata. 


Trees like Macaranga tanarius and Trema spp. are probably quite persistent, 
though the latter stages of this succession have not been observed. Presumably 
seedlings of other limestone elements get established and eventually from the 
dominant vegetation once again, shading out the colonisers. 


4. PHYTOGEOGRAPHY AND SIZE OF THE FLORA 


The total number of species recorded for the limestone flora is 1216 distributed 
in 582 genera and 124 families (This includes the true ferns, Gymnosperms and 
Angiosperms). The introduced elements (as weeds and escapes from cultivation) 
number 39 species or 3.2% ‘of the total flora. 


Malayan Limestone Flora 183 


The breakdown of the flora is as follows: 


Dubious 


Genera Species 
P Records 


PTERIDOPHYTES 

Adiantaceae 8 16 eee 
Dennstaedtiaceae 17 45 = 
Grammitidaceae l - ees 
Hymenophyllaceae l 4 alt 
Polypodiaceae ll 9% ary 
Schizaeaceae 2 3 = 
Thelypteridaceae 3 6 ra! 


TOTAL 


Il 
b 
Ww 
\o 
=i 


GY MNOSPERMS 
Araucariaceae l 
Cycadaceae l 
Gnetaceae l 
Podocarpaceae l 

4 


TOTAL 


HS. aD: me 
| 


Il pe 


ANGIOSPERMS 


DICOTYLEDONS 

Acanthaceae 16 fs 

Amaranthaceae 2 2 l 
l 


w 
NM 


Anacardiaceae 8 
Annonaceae 19 
Apocynaceae 10 
Aquifoliaceae 
Araliaceae 
Aristolochiaceae 
Asclepiadaceae 
Balanophoraceae 
Balsaminaceae 
Begoniaceae 
Bignoniaceae 
Bombacaceae 
Boraginaceae 
Burseraceae 
Buxaceae 
Caprifoliaceae 
Capparidaceae 
Cardiopteridaceae 
Celastraceae 
Chloranthaceae 
Combretaceae 
Compositae 
Connaraceae 
Convolvulaceae 
Cucurbitaceae 
Datiscaceae 
Dilleniaceae 
Dipterocarpaceae 
Ebenaceae 
Elaeocarpaceae 


— WwW 
“IO © 


lon 
| 


nN 


— 


ES eal aati AR ik ee ft, el Seals Seen ha A ties a ee AEN |. a oe 
_—_— — 


184 Gardens’ Bulletin, Singapore — XXX (1977) 


Dicotyledons (cont.) 


Ericaceae 
Erythroxylaceae 
Flacourtiaceac 
Fagaceae 
Eacourtiaceae 
Gentianaceae 
Gesneriaceae 
Guttiferae 
Hamamelidaceae 
Hernandiaceae 
Hypericaceae 
Icacinaceae 
Labiatae 
Lauraceae 
Lecythidaceae 
Leguminosae 
Lentibulariaceae 
Loganiaceae 
Loranthaceae 
Lythraceae 
Malvaceae 
Melastomaceae 
Meliaceae 
Menispermaceae 
Monimiaceae 
Moraceae 
Myristicaceae 
Myrsihaceae 
Myrtaceae 
Nepenthaceae 
Nyctaginaceae 
Ochnaceae 
Oleaceae 
Onagraceae 
Oxalidaceae 
Passifloraceae 
Piperaceae 
Pittosporaceae 
Polygalaceae 
Polygonaceae 
Primulaceae 
Ranunculaceae 
Rhamnaceae 
Rhizophoraceae 
Rosaceae 
Rubiaceae 
Rutaceae 
Santalaceae 
Sapindaceae 
Sapotaceae 
Scrophulariaceae 
Solanaceae 
Staphyleaceae 
Sterculiaceae 
Thymelaeaceae 
Tiliaceae 
Ulmaceae 
Urticaceae 
Verbenaceae 
Violaceae 
Vitaceae 


TOTAL 


Ww 


Om nOON==NhemewUe Tb 
N 


— 


— 


NS) 
— i We Om Oe Ao oo Rh Noe 


NN 


— 


SNR RR KK WK WN KH WK HNK RPRANWHK DAA ot —=ARwWe 
—_ 


nN 
— ON 
We AR OO RANK UNS — NRK RNS = 


— 


CO Ar wmONR— OK NWA RL — CO 
— — 
OhANWIW 


\o 
Ww 
Ww 


805 11 


It 
H 


Malayan Limestone Flora 185 


MONOCOTYLEDONS 


Agavaceae 
Alismataceae 
Amaryllidaceae 
Araceae 
Burmanniaceae 
Commelinaceae 
Cyperaceae 
Dioscoreaceae 
Flagellariaceae 
Gramineae 
Hypoxidaceae 
Liliaceae 
Lowiaceae 
Marantaceae 
Musaceae 
Orchidaceae 
Palmae 
Pandanaceae 
Stemonaceae 
Taccaceae 
Triuridaceae 
Zingiberaceae 


TOTAL 22 


tr —_ 
Ne oe br et eer eer Sip il = 
» 


— (NN 


mn 
—_— 


GRAND TOTAL 126 


A 
les} 
“ 
ON 
a 


The total number of families presented here include Aristolochiaceae and 
Alismataceae which have only been dubiously recorded. Thus the number of 
families actually recorded from the limestone is 124. The total number of plants 
recorded from limestone is 1216 and excluding the ferns this give a total of 1119 
species of spermatophytes. Keng (1970) estimated that between 8000-8500 species 
of spermatophytes occur in the Malay Peninsula. Thus the total number of species 
of spermatophytes on the limestone is between 13.0-13.9% of the total number 
of species found in the Peninsula. 


Henderson (1939) compared the number of species on limestone and the area 
of limestone with the total number of species in Malaya and the total area of 
Malaya. Based on Scrivenor, in Burkill (1935), Henderson concluded that the 
total area of limestone, which is about 260 sq. km. (100 sq. miles), is 0.2-3% of 
the total area of the Peninsula. The total number of species from limestone 
recognised then was 745 which was about 8-9% of the total number of flowering 
plants (then known) in the Peninsula. These figures seem to indicate a much richer 
flora per unit area on the limestone than on the other formations. The present 
figure of 13.0-13.9% would suggest an even richer flora. 


This method of comparison assumes that there is a direct relationship between 
the number of species and the area of land, and that the number of species increases 
linearly with the increase in land area, This seems incorrect. For instance, Gua 
Batu supports about 170 species of plants. This is 14.5% of the total number 
recorded from limestone. It has an area of only about 2.59 sq. km. (1 sq. mile), 
which is 1% of the total area of limestone in Malaya. It would appear that Gua 
Batu were floristically 14.5 times richer than the other limestone hills. 


To obtain a correct picture, only the ‘minimal area’ of a community should 
be used in comparison with the ‘minimal area’ of another community. This 
concept of ‘minimal area’ (Braun-Blanquet, 1932; Poore, 1955) has been defined 


186 Gardens’ Bulletin, Singapore — XXX (1977) 


as the smallest area which can contain an adequate representation of an associa- 
tion. However, the ‘minimal area’ for both the limestone and the lowland forest 
in Malaya have not been worked out. One alternative, therefore, is to select an 
area with the same or about the same area as the limestone and whose flora has 
been worked out. This flora could be used to compare the limestone flora with. 


The flora of Penang island which is about 285 sq. km. (110 sq. mi.) in area 
has been worked out by Curtis (1894). His list of flowering plants total 1805 
species. Similarly the flora of Singapore (Ridley, 1900) which is about 582 sq. km. 
(225 sq. mi.) included 1952 species. The two floras give an idea of the richness 
of the Malayan flora but as far as a comparison with the limestone flora go, they 
do not give an accurate picture. The limestone flora is from an edaphic climax 
formation whereas areas like Singapore and Penang support (originally) a climatic 
climax formation which includes a number of other climaxes, e.g.; mangrove 
swamp forest, beach forest, hill dipterocarp forest (in Penang), Adinandra forest 
(in Singapore), riparian fringes and seasonal swamp forest (at least in Singapore) 
apart from the lowland dipterocarp forest. 


The other alternative therefore is to compare a “‘minimal area”’ of the lime- 
stone flora with the “‘minimal area” of the flora of another edaphic climax forma- 
tion. This comparison will probably give the most accurate picture of the relative 
size and richness of the limestone flora. Unfortunately no flora of any other 
edaphic climax in Malaya has been worked out that could be conviniently used. 


However, Anderson (1963) published an account of the past swamp forest 
of Sarawak and Brunei, which under the present circumstances should be the most 
relevant to compare the limestone flora with. The coastal and deltaic peat swamps 
cover a total of 15644 sq. km. (6040 sq. mi.) of Sarawak and Brunei. The total 
number of flowering plants recorded in Anderson’s list is 393 species. Thus in 
comparison even the vastness of the Sarawak and Brunei peat swamps have not 
support a flora as rich as that on the Malayan limestone. The limestone vegetation 
of Sarawak (surface area much less than the peat swamp) is also exceptionally 
rich in plant species. A preliminary report on the Sarawak limestone (Anderson, 
1965) records over 600 species, and this number is bound to increase as more 
limestone areas are explored. 


Thus one can conclude that the limestone flora of Malaya (and Sarawak) is 
extremely rich in species, intrinsically and also relatively, in comparison with the 
peat swamp flora of Sarawak and Brunei, and most probably with floras of other 
edaphic climaxes. 


The total number of families recorded from limestone is 124; of these 117 are 
families of spermatophytes. The total number of families of flowering plants 
recorded from Malaya (both native and introduced) is 188 (figure obtained by 
totalling the list in Keng, 1969). The delimitation of families in this study is the 
same as in Keng except for Icacinaceae, which is split into Cardiopteridaceae and 
Icacinaceae, and Tiliaceae which has been treated as Elaeocarpaceae and Tiliaceae. 
With the addition of these two families the total number of plant families in Malaya 
becomes 190; the total recorded from limestone amounts to 61.6% of this. 


This rich flora differs from that of the rest of Malaya only in degree and not 
in kind (Henderson 1939). No one family or group of families predominates or is 
confined to the limestone except for two small families represented by very few 
species in Malaya, Buxaceae (three species) and Primulaceae (one species). The 
representation of a family on the limestone also more or less reflects the size of 
the family as represented in Malaya. Table 1 shows the twelve largest families in 
Malaya (from Keng, 1969) which is based on Ridley, 1925, with the total genera 
and species from limestone compared beside them. 


Malayan Limestone Flora 187 


TABLE 1. GENERA AND SPECIES IN THE TWELVE LARGEST (SPECIESWISE) 
FAMILIES IN MALAYA AND THEIR OCCURRENCE ON LIMESTONE 


Families Genera Species 
Malaya Limestone Malaya Limestone 

Orchidaceae 104 52 669 136 
Rubiaceae 67 27 368 66 
Euphorbiaceae 71 37 343 81 
Leguminosae 66 19 266 31 
Gramineae 83 26 205 32 
Annonaceae 30 19 184 39 
Palmae 30 12 181 18 
Melastomaceae 25 8 174 18 
Lauraceae 16 8 174 13 
Acanthaceae 36 16 168 32 
Gesneriaceae 21 11 161 39 
Zingiberaceae 20 9 157 16 
Averages: Ratio z : 1 6 : 1 


On limestone, Orchidaceae still tops the list but there are proportionally more 
members of the Euphorbiaceae than Rubiaceae. The limestone seems a little poor 
in Leguminosae, Palmae, Melastomaceae, Lauraceae and Zingiberaceae though 
the other families are more or less proportionally represented. The reasons for this 
difference may be various and not conjectured as knowledge on the requirements 
of these plants is almost non-existent. 


The families recorded from Malaya but not represented on the limestone are: 


GYMNOSPERMS Te hemtstie eae Simaroubaceae 

. onystylaceae Stylidiaceae 
as 8 aie gn oe pi Shae 

£ Hydrophyllaceae ymplocaceae 
Dicotyledons Illiciaceae Th sdeeak 
Aceraceae Juglandaceae Turneraceae 
Actinidiaceae Linaceae Umbelliferae 
Aizoaceae 2 25 iE ON 
Ancistrocladaceae Malpighiaceae 
Basellaceae Monotropaceae Monocotyledons 
Bixaceae Moringaceae Bromeliaceae 
Cactaceae Myricaceae Butomaceae 
Campanulaceae Nymphaeaceae Cannaceae 
Caricaceae Olacaceae Eriocaulaceae 
Caryophyllaceae Opiliaceae Hydrocharitaceae 
Casuarinaceae Orobanchaceae Iridaceae 
Chenopodiaceae Pedaliaceae Juncaceae 
Clethraceae Plantaginaceae Lemnaceae 
Cornaceae Plumbaginaceae Najadaceae 
Crassulaceae Portulacaceae Philydraceae 
Cruciferae Proteaceae Pontamogetonaceae 
Cunoniaceae Rafflesiaceae Pontederiaceae 
Daphniphyllaceae Sabiaceae Restionaceae 
Dichapetalaceae Salicaceae Smilacaceae 
Droseraceae Saxifragaceae Typhaceae 
Elaeagnaceae Schisandraceae Xyridaceae 


These 71 families can be divided into six groups. 


1. Aquatic or marsh plants. These are represented by 12 families, and include 
most of the monocots. They are, Nymphaeaceae, Stylidiaceae, Butomaceae, 
Eriocaulaceae, Hydrocharitaceae, Lemnaceae, Najadaceae, Philydraceae, Ponta- 
mogetonaceae, Pontederiaceae, Restionaceae, and Typhaceae. 


188 Gardens’ Bulletin, Singapore — XXX (1977) 


2. Parasitic plants. These are represented by two families, Orobanchaceae 
and Rafflesiaceae. 


3. Coastal plants. Represented by three families, Casuarinaceae, Goodenia- 
ceae and Portulacaceae. Aizoaceae also has a coastal member in Sesuvium, while 
the remaining two genera in Malaya are not very common herbs. 


4. Weeds or plants in cultivation. These are represented by 17 families, 
Pinaceae, Basellaceae, Bixaceae, Caricaceae, Caryophyllaceae, Cactaceae, Cheno- 
podiaceae, Crassulaceae, Cruciferae, Moringaceae, Pedaliaceae, Plantaginaceae, 
Plumbaginaceae, Salicaceae, Turneraceae, Bromeliaceae and Iridaceae. 


5. Very rare plants. In many cases these families are represented by a 
solitary species in Malaya. There are 15 such families. They are, Aceraceae, 
Ancistrocladaceae, Clethraceae, Cunoniaceae, Daphniphyllaceae, Droseraceae, 
Elaeagnaceae, Epacridaceae, Hydrophyllaceae, Illiciaceae, Magnoliaceae, Mono- 
tropaceae, Myricaceae, Cannaceae and Juncaceae. 


6. Others. These remaining 23 families are often small and uncommon. They 
are Actinidiaceae, Aizoaceae, Campanulaceae, Cornaceae, Dichapetalaceae, Gony- 
stylaceae, Juglandaceae, Linaceae, Malpighiaceae, Olacaceae, Opiliaceae, Protea- 
ceae, Sabiaceae, Saxifragaceae, Schisandraceae, Simaroubaceae, Styracaceae, 
Symplocaceae, Theaceae, Umbelliferae, Smilacaceae and Xyridaceae. 


It is unlikely that members of group 1 will ever be found on limestone, but 
it is probable that members of group 2 are present on limestone. Members of 
group 3 could conceivably be present on coastal limestone barring any physiological 
antagonism to limestone. When more limestone areas are disturbed by man, some, 
particularly the weed members of group 4 will probably be recorded. Members 
group 5 which are very rare plants will, even if they are recorded on limestone, 
also be very rare. New records for limestone from families not yet represented will 
therefore most likely be from group 6, but there will not be many species, for most 
of these families are small. The greatest number of new records will probably be 
from families already represented on the limestone. 


Phytogeography 


The 1216 species recorded from the limestone can be grouped according to 
their geographical distribution into a number of classes, viz: 


PANTROPIC: Species occurring throughout the tropical regions of the world. 


PALEOTROPIC: Species occurring in the tropics of the Old World, from Africa 
and India through Malesia and to the Pacific. 


INDO-MALESIAN: Species disturbed in mainland Asia as well as in Malesia; 
including India to the Himalayas, Burma, S. China, Indochina, Thailand, 
Malesia and southwards to Australia and the Pacific Islands. 


ASIATIC: Species only in mainland Asia, sometimes only from Thailand. 


MALESIAN: Species occurring in Malesia, including Malaysia, Indonesia, — 


Philippines, New Guinea and sometimes to Australia and the Pacific. 


ENDEMIC: Species occurring only in Malaya. These are subdivided into 
(a) Species restricted to limestone in Malaya. 
(b) Species not restricted to limestone in Malaya. 


Malayan Limestone Flora 189 


The result of this analysis is shown in Table 2. 


TABLE 2. GEOGRAPHICAL DISTRIBUTION OF THE SPECIES 
NUMBER OF SPECIES 


Pterido- Gymno- Angio- 

phytes sperms sperms Total To 
PANTROPIC 6 — 14 20 1.7 
PALEOTROPIC 8 --- 27 35 29 
INDO-MALESIAN 40 4 315 359 29.6 
ASIATIC (Asia mainland) 5 1 228 234 19.0 
MALESIAN 32 1 225 257 21.2 
ENDEMIC (total) 5 — 266 272 22.5 
(a. restricted to limestone 2 — 127 129 10.7) 
(b. not restricted to limestone 3 — 139 143 11.8) 
WEEDS AND ESCAPES 1 —_ 38 39 2 fd 


From the table it can be seen that the pantropic element is negligible; this is 
because most species of this group are weeds (plants that occur in cultivated 
habitats as undesirables, also in lawns and other open places and often as members 
of secondary vegetation) and have been included under that group. The widely 
distributed paleotropic element is also small. There is no predominance of either 
the Asiatic or Malesian element, both being equally represented except for the 
ferns. The weeds and escapes are noticably few in number; this is probably because 
there are few areas on the limestone that support a secondary vegetation. However, 
this is not to say that the limestone hills have been free from interference, but, that 
when the limestone hills are disturbed it usually means that the habitat itself is 
bodily removed as for instance in mining and quarrying. 


Krasan (1882) and Steenis (1934) suggested that on limestone mountains 
plants might be able to descend abnormally low. Although there are no really high 
limestone mountains in Malaya this tendency can be seen. Here the mountain 
massif has many peaks rising to over 1500 m, with Gunong Tahan rising to 2187 m. 
Plants known from 800-2000 m (very rarely from lower elevation, if ever) have 
been ae on low elevation (usually jess than 300 m) on the limestone and not 
elsewhere. 


The species recorded include Antrophyum  semicostatum, Leptochilus 
decurrens, Vittaria angustifolia, Alyxia angustifolia, Alyxia pumila, Corybas, 
mucronatus, Distylium stellare, Liparis compressa, Paphiopedilum lowii, Schefflera 
elegans, Sciaphila asterias, Scycopsis dunnii and Toxocarpus curtisii. 


5. AFFINITIES OF THE FLORISTIC COMPONENTS TO LIMESTONE 


The component species of the Malayan limestone can be arranged in the 
following four groups: 


I. EXCLUSIVES: These are species which, in Malaya, are restricted to the 
limestone and include, (a) Species endemic to Malaya. (b) Species not 
endemic to Malaya. 


II. PREFERENTS: Species with a preference for the limestone field, appear- 
ing in both fields (limestone and non-limestone) but more abundantly in the 
limestone field. 


Ill. INDIFFERENTS: Species with no affinity for either field, appearing in 
both fields without exhibiting any difference in abundance between them. 
This group includes most epiphytes and plants of secondary vegetation. 


IV. STRANGERS: Species appearing accidentally on the limestone field; 
frequently collected from the non-limestone field but only rarely from the 
limestone field. 


The criterion of fidelity (Brown-Blanquet, 1932) is used in constructing this 


grouping, fidelity being indicative of the degree with which a species is restricted 
to a particular kind of community (in this case the limestone community). 


190 Gardens’ Bulletin, Singapore — XXX (1977) 
The fidelity of each species and thus the allocation to the various groups is 
based solely on field work. Species under groups I and II are termed the 
characteristic species. 

Rarely collected species which have been recorded both from limestone and 
non-limestone fields are included under group II (indifferents) instead of under 
group IV (strangers). This is because the limestone areas are probably the least 
botanised of all the vegetation types in Malaya and chances are that rare species 
already recorded from limestone are more likely to turn up from it again rather 
than from the other vegetation types. 


Weeds and plants escaped from cultivation usually fit into groups III (indiffe- 
rents) or IV (strangers). However, here they are listed separately. 

The true and intrinsic reasons for a plant to be on the limestone field could 
also be used to classify the floristic components. Any of the plants found on 
limestone could be: 


(a) Calciphilous (chalk-loving) in the true physiological sense (Jackson, 1928). 
They may be obligate calciphiles, in which case the limestone field is a prerequisite 
for growth and survival, or non-obligate. The non-obligates have a physiological 
need or affinity for limestone but are able to survive without the limestone field. 

A large number of species in my group I (exclusives) are probably obligate 
calciphiles while some (those that are non-endemic which are also found away 
from the limestone outside Malaya) are probably non-obligate calciphiles. These 
non-obligate calciphiles will also include a large number of species from my 
group II (preferents). 


(b) Indifferents; species that are able to tolerate or are unaffected, physiologically, 
by the limestone habitat; as in my group III. 


(c) Calcifugal (shunning chalk) in the true physiological sense (Jackson, (1928). 
Obligate calcifuges cannot tolerate the limestone field whereas the non-obligate 
calcifuges may often appear as strangers on the limestone field when the other 
criteria of space, competition and moisture are favourable. The non-obligates 


include at least some of my group IV. 
However, this classification can only be effected by experimental work. 
The species list of the different groups follows and are summed up in Table 3. 


PTERIDOPHYTES 


Adiantum zollingeri 
Antrophyum parvulum 
Cheilanthes farinosa 
*Doryopteris allenae 
Doryopteris ludens 
Drynaria bonii 
Heterogonium alderwereltii 
Heterogonium pinnatum 
Lygodium polystachyum 
Nephrolepis dicksonioides 
*Polystichum lindsaeifolium 
Pyrrosia stigmosa 
Schizaea inopinata 
Tectaria amplifolia 
Tectaria devexa 


GYMNOSPERMS 
Cycas siamensis 


ANGIOSPERMS 
DICOTYLEDONS 


Actephila excelsa 
Aeschynanthus longicaulis 


I. EXCLUSIVES 


*Amaracarpus saxicola 

Andrographis tenuiflora 

*Aporuellia sumatrensis var. 
ridleyi 

*Adisia biflora 

*Ardisia langkawiensis 

*Ardisia meziana 

Argostemma diversifolium 

*Barleria siamensis var. 
glabrescens 

Begonia curtisii 

*Begonia foxworthyi 

*Begonia ignorata 

*Begonia kingiana 

*Begonia nurii 

*Begonia phoeniogramma 

Boea acutifolia 

*Boea brachycarpa 

*Boea caerulescens 

*Boea divaricata 

*Boea lanata 

*Boea minutiflora 

*Boea paniculata 

*Boea parviflora 

*Boea suffruticosa 


*Endemic to Malaya. 


Boea treubii 

*Boea verticillata 
Boerhavia chinensis 
*Buxus holttumiana 
*Buxus malayana 
*Buxus rupicola 
*Callicarpa angustifolia 
*Canarium perlisanum 
Canscora pentanthera 
*Canthium aciculatum 
Capparis diffusa 
Celtis philippensis 
*Chirita caliginosa 
Chirita hamosa 
Chirita involucrata 
Chirita lacunosa 
*Chirita rupestris 
*Chirita sericea 
*Cleistanthus parvifolius 
Cnesmone laevis 
*Cnesmone subpeltata 
Colona javanica 
Cynoctonum mitreola 
Dendronide sinuata 
Dichiloboea speciosa 


Malayan Limestone Flora 


Dicotyledons (cont.) 


*Dicliptera rosea 
Dimocarpus longan ssp. 
longan var. longan 

Diospyros daemona 
*Diospyros holttumii 
Diospyros retrofracta 
Diospyros transitoria 
Diospyros undulata 
*Dischidia tomentella 
*Drypetes nervosa 
*Embelia calcarea 
Euphorbia antiquorum 
Excoecaria oppositifolia 
*Fagraea calcarea 
Ficus calcicola 
Garuga floribunda 
Glossocarya mollis 
*Glycosmis calcicola 
*Glycosmis calcicola var. 
kelantanica 
Glyptopetalum zeylanicum 
*Gongylosperma lanuginosum 
Goniothalamus subevenius 
Grewia viminea 
Gymnanthera insularum 
*Gymnostachyum robinsonii 
Heritiera pterospermoides 
*Homalium kunstleri 
Homalium undulatum 
*Hoya occlusa 
Hydnocarpus ilicifolia 
*Impatiens crytoneura 
Impatiens marosepala 
Impatiens mirabilis 
Impatiens opinata 
*Impatiens ridleyi 
Impatiens scortechinii 
*Impatiens tipusensis 
Impatiens vaughanii 
*Isonandra perakensis var. 
kelantanensis 
*Isonandra perakensis var. 
perakensis 
*Jasminum cordatum 
*Jasminum curtisii 
*Jasminum sp. A aff. 
trinerve 
*Jasminum sp. B 
*Justicia microcarpa 
*Justicia robinsonili 
*Justicia rupestris 
*Justicia subalternans 
Justicia valida 
*Kopsia griffithii var. 
paucinervia 
*Lagerstroemia langkawiensis 
*Lasiobema flavum 
Leptopus australis 
Leucas mollissima 
Ligustrum confusum 
Lysimachia peduncularis 
*Madhuca calcicola 
*Mallotus bracteatus 
Mallotus brevipetiolatus 
Mallotus cuneatus 
Maytenus curtisii 
*Melodinus perakensis 
Microphium pubescens 


*Miliusa parviflora 
*Millettia pterocarpa 
Monophyllaea glabra 
*Monophyllaea hirticalyx 
Oidentandia rosettifolia 
*Ophiorrhiza fruticosa 
Ophiorrhiza kunstleri 
*Ophiorrhiza longerepens 
Ornithoboea flexuosa 
Orophea cuneiformis 
*Orophea hirsuta 
Orophea polycarpa 
*Osmanthus scortechinii 
*Paraboea bakeri 
*Paraboea bettiana 
*Paraboea ferruginea 
*Paraboea laxa 
*Paraboea vulpina 
Parishia rosea 
*Pavetta pauciflora 
Pentacme siamensis 
Pentaspadon curtisii 
Peperomia dindigulensis 
Peperomia portulacoides 
*Peperomia sp. A. 
*Phanera decumbens 
Phyllanthus columnaris 
Phyllanthus ridleyanus 
*Pilea fruticosa 
*Piper collinum 

Pisonia aculeata 
Piscnia umbellifera 
*Pistacia malayana 
*Plectranthus kunstleri 
Polygala cardiocarpa 
Polygala malesiana 
Polygala triphylla 
*Polytrema cupreum 
*Popowia velutina 
*Rungia minutiflora 
Sageretia thea 

Sapium insigne 
*Sauropus calcareus 
Sauropus macranthus 
Sauropus suberosus 
*Scheffera subracemosa 


Scleropyrum wallichianum 


*Scolopia steenisiana 
Semecarpus glomerulata 
Solanum biflorum 
*Sonerila elliptica 
Sonerila tenera 
*Stenothyrsus ridleyi 
Stephania venosa 
Sterculia angustifolia 
Sterculia lancaviensis 


*Strobilanthes pachyphyllus 


Tarenna angustifolia 
*Tarenna calcarea 
Tarenna curtisii 
Tetrameles nudiflora 
Timonius atropurpureus 
*Toxocarpus pauciflorus 


Trigonostemon verticillatus 


*Tylophora calcicola 
Vernonia curtisii 
*Vernonia rupicola 
Vitex siamica 


191 


*Vitis scortechinii 
Wikstroemia 
androsaemifolia 
*Zanonia clarkei 
*Zizyphus pernettyoides 


MONOCOTYLEDONS 


*Aglaonema costatum 
Amorphophallus carnosus 
Amorphophallus 

haematospadix 

Arisaema fimbriatum 
Arthraxon prionodes 
Asparagus racemosus 
*Boesenbergia curtisii 
*Calamus balingensis 
Calanthe rubens 
Calanthe vestita 
*Carex malaccensis 
Carex speciosa 

*Corymborchis brevistylis 
Cymbopogon calcicola 
*Dendrobium langkawiense 
Dendrocalamus dumosus 
*Dendrocalamus elegans 
Dichanthium annulatum 
*Dichanthium mucronulatum 
Dracaena curtisii 
Dracaena yuccaefolia 
Eulalia quadrinervis 
Eulophia keithii 
*Fimbristylis calcicola 
*Fimbristylis malayana 

*Fimbristylis trichophylla 
*Globba albiflora var. aurea 
Goodyera hispida 
Habenaria carnea 

*Habenaria kingii 
Hapaline brownii 
*Isachne langkawiensis 
Kaempferia elegans 
Kaempferia pulchra 
*Liberbaileya gracilis 
*Malaxis reniloba 
*Malleola undulata 
*Maxburretia rupicola 

*Oberonia calcicola 
Oberonia caudata 

*Oberonia transversiloba 
*Pandanus calcicola 
*Pandanus irregularis 
*Pandanus piniformis 
Paphiopedilum niveum 
Pollia subumbellata 
Pomatocalpa naevatum 
Pomatocalpa setulense 

*Pothos lorispatha 

*Pteroceras tanyphyllum 

*Raphidophora kunstleri 
Sarcanthus termissus 
Spathoglottis hardingiana 

*Stachyphrynium cylindricum 

*Thelasis succosa 
Trichoglottis winkleri var. 

minor 

*Typhonium filiforme 

*Typhonium fultum 


*Endemic to Malaya. 


192 


PTERODOPHYTES 


Adiantum malesianum 
Adiantum soboliferum 
Asplenium adiantoides 
Asplenium squamulatum 
Cyclopeltis crenata 
Hemionitis arifolia 
Pteridrys syrmatica 
*Pteris longipinnula var. b 
Pyrrosia penangiana 


GYMNOSPERMS 
ANGIOSPERMS 


DICOTYLEDONS 


*Argostemma inaequilaterum 
Argostemma pictum 
Berrya cordifolia 
Chirita viola 
Cladogynos orientalis 
Cleistanthus gracilis 
*Cleistanthus kingii 
Colona merguensis 
Croton cascarilloides 
Cymaria dichotoma 
*Debregeasia squamata 
Deeringia polysperma 
*Dehaasia curtisii 
esmodium rugosum 


PTERIDOPHYTES 


Adiantum stenochlamys 
Asplenium macrophyllum 
Asplenium pellucidum 
Asplenium salignum 
Davallia denticulata 
Davallia solida 
Drynaria quercifolia 
Drynaria rigidula 
Humata pectinata 
Loxogramme avenia 
Loxogramme scolopendrina 
Nephrolepis falcata 
Photinopteris speciosa 
Phymatodes nigrescens 
Phymatodes papillosum 
Phymatodes scolopendria 
Pityrogramma calomelanos 
Pteridium aquilinum var. 
wightianum 
Pteridium caudatum var. 
yarrabense 
Pteris ensiformis 
Pteris tripartita 
Pteris vittata 
Pyrrosia adnascens 
Pyrrosia varia 
Tectaria variolosum 
Thelypteris immersa 
Trichomanes bipunctatum 


Gardens’ Bulletin, Singapore — XXX (1977) 


Il. PREFERENTS 


*Drypetes oxyodonta 
Epithema saxatile 
Fagraea carnosa 
Fagraea curtisil 

Ficus curtipes 

Garcinia minutiflora 
Gomphostemma crinitum 
Hedyotis coronaria 
*Heterostemma piperifolium 
Hopea ferrea 

*Hoya citrina 

Illigera pulchra 
*Ixora clerodendron 
Jasminum wrayi 
*Justicia henicophylla 
Lasiobema curtisii 
*Lasiobema strychnoideum 
*Leea saxatilis 

Litsea glutinosa 
*Madhuca ridleyi 
Mallotus dispar 
Mallotus miquelianus 
Memecylon laevigatum 
*Miliusa amplexicaulis 
Monophyllaea horsfieldii 
Murraya paniculata 
*Orophea maculata 
*Paraboea capitata 
*Peperomia kotana 
*Phyllanthus filicifolius 
Procris pedunculata 


il. INDIFFERENTS 


GYMNOSPERMS 


Cycas rumphii 
Podocarpus polstachyus 


ANGIOSPERMS 


DICOTYLEDONS 


Actephila ovalis 

Adenia nicobarica 
Aeschynanthus parvifolia 
*Aglaia argentea 

Aglaia odoratissima 

Aglaia splendens 
Agrostistachys gaudichaudii 


Allophylus cobbe var. glaber 


Allophylus cobbe var. 
villosus 
Alyxia selangorica 
Antidesma japonicum 
*Ardisia fulva 
*Ardisia kunstleri 
Ardisia lancelolata 
Ardisia oxyphylla 
*Ardisia solanacea var. elata 
*Ardisia vaughani 
Argyreia maingayi 
Argyreia mollis 
*Artabotrys grandifolius 
Atalantia monophylla 
*Atalantia roxburghiana 


Rinorea bengalensis 
Sauropus villosus 
Sumbaviopsis albicans 
Terminalia triptera 
Trigonostemon viridissimus 
Vitis discolor 

Wikstroemia indica 
Zizyphus oenoplia 


MONOCOTYLEDONS 


Abdominea minimiflora 
Amomum testaceum 
Apluda mutica 
Appendicula torta 
Arachnis flos-aeris 
Bulbophyllum fenestratum 
*Bulbophyllum flammuliferum 
Bulbophyllum lilacinum 
Dendrobium salaccense 
Dioscorea calcicola 
Dracaena congesta 
*Dracaena graminifolia 
*Globba fasciata 
Habenaria reflexa 
Pholidota pallida 
*Pothos macrocephalus 
*Sarcanthus rugulosus 
Trichoglottis retusa 
Uncifera tenuicaulis 


Becheria parviflora 
*Begonia debilis 

Begonia guttata 
*Beilschmiedia lumutensis 
Bombax anceps 

Bridelia ovata 

Bridelia tomentosa 
Callicarpa lanata 
Canthium didymum 
Carallia brachiata 
Casearia capitellata 
Cassia timoriensis 
Cinnamomum iners 
Clausena excavata 
*Cleidion javanicum 
Cleistanthus decurrens 
*Cleistanthus glaucus 
*Cleistanthus macrophyllus 
Clerodendron penduliflorum 
Clerodendron serratum 
Clidemia hirta 
Combretum porterianum 
Congea vestita 
Connarus sp. 

Cordia griffithii 

Cordia obliqua 
Cratoxylum maingayi 
Cyclea laxiflora 
*Dalbergia kunstleri 
Dalbergia phyllanthoides 
Dalbergia scortechinii 


*Endemic to Malaya. 


Malayan Limestone Flora 


Dicotyledons (cont.) 


Decaspermum fruticosum 
*Dehaasia longipedicellata 
Dendrocnide stimulans 
Desmos cochinchinensis 
Desmos dasymaschalus var. 
wallichii 
Desmos dunalii 
*Diospyros adenophora 
Diospyros buxifolia 
Diospyros cauliflora 
*Diospyros ellipsoidea 
Diospyros ferrea 
Diospyros frutescens 
Diospyros toposiodes 
Dischidia benghalensis 
Dischidia hirsuta 
*Dischidia scortechinii 
Ehretia timorensis 
*Elatostema curtisii 
Elatostema latifolium 
*Enicosanthum congregatum 
Eriobotrya bengalensis 
Erismanthus obliquus 
*Ervatamia peduncularis 
Erythroxylum cuneatum 
*Eugenia pendens 
*Eugenia porphyranthera 
Euonymus cochinchinensis 
Fagraea blumei 
Ficus binnendykii 
Ficus deltoidea 
Ficus elastica 
Ficus hispida 
Ficus microcarpa 
Ficus sagittata 
Ficus subulata 
Ficus sundaica 
Ficus superba var. japonica 
Ficus tinctoria ssp. gibbosa 
Ficus virens var. glabella 
Ficus villosa 
Garcinia murdochii | 
Garcinia nigrolineata 
*Garcinia opaca 
Geophila repens 
Glochidion rubrum 
Glycosmis puberula 
Glycosmis rupestris 
*Glycomis sapindoides 
Glytopetalum quadrangulare 
Gomphia serrata 
Gomphostemma javanicum 
Gomphostemma microcalyx 
*Goniothalamus fulvus 
Grewia acuminata 
*Gymnostachyum decurrens 
*Gymnostachyum 
diversifolium 
Hedvotis tenellifiora 
Hedyotis verticillata 
Helicteres angustifolia 
Helicteres hirsuta 
Helixanthera axillaris 
Helixanthera pulchra 
Hemigraphis ridleyi 
Heritiera littoralis 
Holarrhena curtisii 


Homalanthus populneus © 
Homalium dasyanthum 
Hoya coronaria 

Hoya latifolia 

*Hoya maingayi 

Hoya parviliora 

Hoya ridleyi 
Hydnocarpus wrayl 
Iodes cirrhosa 

Iodes ovalis 

Ipomoea illustris 

Ixora nigricans var. ovalis 
Ixora pendula 

Ixora umbellatta var. 

multibracteata 

Jacquemontia paniculata 
Jasminum adenophyllum 
Jasminum insularum 
*Justicia pectinella 
*Justicia ptychostoma 
*Justicia subcymosa 
*Justicia uber 

Knema globularia 
Knema laurina 

Kopsia pauciflora 
Laportea interrupta 
Leea aequata 

Leea rubra 

Leea sambucina 
Leptonychia glabra 
Ligustrum robustum 
Litsea norohae 
Loeseneriella pauciflora 
Macaranga tanarius 
Macrosolen cochinchinensis 
*Maesa pahangiana 
Malaisia scandens 
Mallotus wrayi 
Mammea brevipes 
Marsdenia tinctoria 
*Medinilla scortechinii 
Meiogyne virgata 
Melochia umbellata 
*Melodinus orientalis 
Melothria affinis 
*Memecylon dichotomum 
Memecylon edule 
Memecylon floribundum 
*Memecylon kunstleri 
Memecylon pauciflorum 
*Memecylon wallichii 
Microdesmis casearifolia 
Micromelum minutum 
Milusa longipes 
Mimusops elengi 
Momordica subangulata 
Morinda elliptica 
Morinda umbellata 
Mucuna biplicata 
Mycetia malayana 
Myrsine porteriana 
Naravelia dasyoneura 
Neolitsea zeylanica 
Oldenlandia ovatifolia 
Oldenlandia pterita 
Ophiorrhiza discolor 
Ophiorrhiza hispidula 


*Endemic to Malaya. 


193 


*Ophiorrhiza pallidula 
Ophiorrhiza remotiflora 
Orophea enterocarpa 
Orthosiphon aristatus 
Oxymitra biglandulosa 
Pachycentria constricta 
Paederia tomentosa 
Paramignya scandens 
Paranephelium 

macrophyllum 
Payena lucida 
Petunga hirta 
Phanera glauca 
Phyllanthus oxyphyllus 
Phyllanthus pulcher 
Phyllanthus sikkimensis 

*Piper mucronatum 

*Piper scortechinii 
Piper umbellatum 
Planchonella obovata 

“Plethiandra sessiliflora 
Pogonanthera pulverulenta 
Poikilospermum suaveolens 

*Polyalthia brunneifolia 

*Polyalthia lateritia 
Polytrema uber 

*Prema rubens 
Prismatomeris malayana 
Pseuderanthemum 

crenulatum 
Pseuderanthemum 

graciliflorum 
Pseuduvaria macrophylla 
Pseuduvaria setosa 

Psychotria angulata 

*Psychotria cantleyi 
Psychotria montana 
Psychotria rhinocerotis 
Pterolobium densiflorum 
Pterospermum jackianum 
Pterospermum pectiniforme 

*Quisqualis parvifolia 
Radermachera lobbii 
Randia densiflora 
Rauwolfia reflexa 

*Richeriella malayana 
Rinorea horneri 
Rostellaria procumbens 

*Rubus angulosus 
Ruellia repens 

*Rhus perakensis 
Salacia grandiflora 
Salacia korthalsiana 
Salacia macrophylla 
Salomonia ciliata 
Saraca declinata 
Saraca thaipingensis 
Secamone micrantha 
Schefflera junghuhniana 

*Schefflera musangensis 
Schefflera subulata 
Schefflera tomentosa 
Schefflera venolusa 
Scolopia spinosa 
Scurrula ferruginea 
Scutellaria discolor var. 

discolor 


194 


Dicotyledons (cont.) 


Sida javensis 
Stauraninera grandifolia 
Steiecnocarpus Caulllorus 
Sverculia rubiginosa 
Suveoius asper 

Sctreoius iuicifolius 
Sireoius (axoides 
*Strobilanthes leucopogon 
Sirycnnos axillaris 
*jarenna puichra 
*Jarenna ridieyl 
‘Lermunatia Calamansanai 
*|oxocarpus Curtisi 
Treina tomentosa 
Trigonostemon aurantiacus 
Trichosanthes tricuspidata 
*Trigonosteinon villosus 
*Tristania subauriculata 
Trivaivaria macrophylla 
Turpinia ovalifolia 
Tylophora perakensis 
Tylophora tenuis 
Vaccinium littoreum 
Vatica cinerea 

Ventilago gladiata 
Ventilago oblongifolia 
Villebrunea sylvatica 
Viscum orientale 

Vitis furcata 

Vitis glaberrima 

Vitis hastata 

*Vitis kunstleri 

*Vitis mollissima 

Vitis pyrrhodasys 

Vitis repens 

*Vitis wrayi 

Wrightia dubia 
Wrightia laevis 


MONOCOTYLEDONS 


*Adenoncos major 
Adenoncos parviflora 
Adenoncos sumatrana 
Aglaonzma oblongifolium 
Agrostophyllum 
bicuspidatum 
Alocasia denudata 
Alocasia lowii 


PTERIDOPHYTES 


Abacopteris urophylla 
Antrophyum callifolium 
Antrophyum semicostatum 
Arcypteris irregularis 
Asplenium phyllitidis 
Asplenium unilaterale 
Athyrium cordifolium 
Athyrium esculentum 
Athyrium montanum 


Gardens’ Bulletin, Singapore — XXX (1977) 


Amorphophallus prainii 
Amorphopnalius variabilis 
Auyariuin hurmile 
Anauvendrum marginatum 
*Arenga hookeriana 
*Arenga weslermoutil 
Afrisaeuia roxburghil 
borassogendron suachadonis 
Bulvvpiylium apodum 
Bulbophyllum concinnum 
Bbuloopnylium 
Meuibranaceum 
Buriuiannia championii 
Burmannia lutescens 
Calamus concinnus 
Calamus siamensis var. 
mailaianus 
Calanthe ceciliae 
Caianthe triplicata 
Camarotis apiculata 
Carex breviscapa — 
Caryota mitis 
Catiunbium speciosum 
Centhotheca lappacea 
Ceratostylis pendula 
Chlorophytum orchidastrum 
Chrysopogon fulvus 
Chrysopogon orientalis 
Coelogyne asperata 
*Coelogyne pallens 
Coelogyne pandurata 
Colocasia gigantea 
Corybas mucronatus 
Corymborchis veratrifolia 
Cymbidium finlaysonianum 
Dendrobium aloifolium 
Dendrobium spurium 
Dichanthium caricosum 
Dioscorea bulbifera 
Dioscorea esculenta 
Dioscorea filiformis 
Dioscorea glabra 
Dioscorea prazeri 
Dracaena angustifolia 
*Dracaena nutans 
Ephemerantha luxurians 
Epipremnum giganteum 
Eria leptocarpa 


IV. STRANGERS 


Athyrium pinnatum 
*Athyrium prescottianum 
Blechnum finlaysonianum 
Crypsinus enervis 
Ctenopteris alata 
Ctenopteris moultoni 
Cyclosorus extensus 
Cyclosorus interruptus 
Cyclosorus megaphyllus 
Cyclosorus unitus 


*Endemic to Malaya. 


Eria pulchella 
Fimbristylis fusca 
Fiubristyius fuscoides 
*Forrestia monosperina 
Hanguana malayana 


*Hoimalomena deitoidea 


*iguanura polyimorpna 
Lasia aculeata 

Liparis caespitosa 
Liparis gibbosa 

Malaxis calophylla 
Matieola dentitera 
Microsaccus ampullaceus 
Microsaccus brevifolius 
Microsaccus javensis 
Neyraudia reynaudiana 
*Oberonia flava 
Oplismenus compositus 
*Pandanus alticola 
*Peliosanthes lurida 
Phalaenopsis cornu-cervi 
Phalaenopsis decumbens 
Phreatia secunda 
Podochilus lucescens 
Podochilus microphyllus 
Pogonatherum paniceum 
Pollia sumatrana 
Polystachya flavescens 
Pomatocalpa spicatum 
Pothos scandens 
Pteroceras ciliatum 
Pteroceras hirsutum 
Renanthera histrionica 
Rhychelytrum repens 
Sarcanthus sacculatus 
Sarcanthus scortechinii 
Sarcanthus subulatus 
Schismatoglottis calyptrata 
*Schismatoglottis mutata 
*Sciaphila asterias 
Scindapsus hederaceus 
Scindapsus perakensis 
Scleria lithosperma 
Stemona tuberosa 
Thelasis micrantha 
Thelasis triptera 
*Tupistra grandis 
Vandopsis gigantea 


Drynaria sparsisora 
Humata heterophylla 
Lemmaphyllum accedens 
Lepisorus longifolius 
Leptochilus decurrens 
Lindsaya lucida 
Lygodium flexuosum 
Microlepia speluncae 
Microsorium musifolium 
Microsorium punctatum 


Malayan Limestone Flora 


Pteridophytes (cont.) 


Nephrolepis biserrata 
Nephrolepis hirsutula 
Nephrolepis radicans 
Oleandra undulata 
Pteris mertensioides 
*Pteris scabripes 
Pyrrosia floccigera 
Taenitis blechnoides 
Tectaria barberi 
Tectaria griffithii 
Tectaria macronta 
Trichomanes christii 
Trichomanes humile 
Trichomanes motleyi 
Vittaria angustifolia 
Vittaria elongata var. 
angustifolia 


GYMNOSPERMS 


Agathis dammara 
Gnetum cuspidatum 
Gnetum gnemon var. 
tenerum 
Podocarpus neriifolius 


ANGIOSPERMS 


DICOTYLEDONS 


Acrotrema costatum 
Adenosma capitatum 
Aeschynanthus radicans 
Agelaea borneensis 
Alchornea rugosa 
Alsomitra pubigera 
Alstonia scholaris 
*Alyxia angustifolia 
*Alyxia pumila 
Anaxagorea javanica 
Antidesma montanum 
Antidesma tomentosum 
*Aporosa stellifera 
Aralidium pinnatifidum 
Ardisia andamanica 
Ardisia crenata 
Ardisia colorata var. 
complanata 
Ardisia pendula 
*Ardisia playtyclada 
Ardisia ridleyi 
*Ardisia tahanica 
Ardisia villosa 
Azadirachta excelsa 
Baccaurea lanceolata 
Balanophora fungosa 
Barleria prionitis 
Barringtonia asiatica 
*Barringtonia fusiformis 
Barringtonia macrostachya 
Bauhinia acuminata 
*Beilschmiedia pahangensis 
Biophytum adiantoides 
Blumeodendron kurzii 
Brassaiopsis polyacantha 
Breynia vitis-idaea 
Bridelia stipularis 
Buchanania sessilifolia 
Caesalpinia crista 


Calophyllum curtisii 
Cananga odorata 
Canarium pilosum 


195 


Ficus trichocarpa var. 
obtusa 
Firmiana malayana 


Canarium pseudodecumanum Garcinia cowa 


Capparis pubiflora 
Cardiopteris javanica 
Casearia grewiaefolia 
Centranthera hispida 
Chloranthus elatior 
Chukrassia tabularis 
Citrus macroptera 
Claoxylon longifolium 
Cleistanthus hirsutulus 
Cleistanthus polyphyllus 
Clerodendron paniculatum 
Cnesmone javanica 
Coffea canephora 
*Coffea malayana 

Coleus scutellarioides 
Colubrina asiatica 
Combretum latifolium 
Connarus semidecandrus 
Cotylelobium malayanum 
Croton argyratus 
Croton erythrostachys 
Croton laevifolius 
Cryptocarya griffithiana 
Curanga amara 
Cynometra malaccensis 
*Cyrtandra cupulata 
*Cyrtandra lanceolata 
*Dacryodes kingii 
Dehassia microcarpa 
Derris elliptica 

Derris thyrsiflora 
Desmodium umbellatum 
Dillenia indica 
Diospyros hermaphroditica 
Diospyros malayana 
Diospyros pilosanthera 
Diospyros rigida 
Diospyros wallichii 
Diploclisia glaucescens 
Dipterocarpus oblongifolius 
Dischidia rafflesiana 
Distylium stellare 
Dryobalanops aromatica 
Dryobalanops oblongifolia 
Dysoxylum arborescens 
Elaeocarpus pedunculatus 
Endospermum diadenum 
Erythrina variegata 
Eugenia chlorantha 
Eugenia claviflora 
Eugenia spicata 
Euonymus javanicus 
Exacum tetragonum 
Fagraea auriculata 
Fagraea ceilanica 

Ficus annulata 

Ficus botryocarpa 

Ficus montana 

Ficus oligodon 

Ficus parietalis 

Ficus racemosa 

Ficus scortechinii 

Ficus semicordata 

Ficus stricta 


*Endemic to Malaya. 


*Garcinia eugeniaefolia 
Garcinia merguensis 
*Garcinia montana 
Glochidion obscurum 
Glochidion perakense 
Glycosmis chlorosperma 
Gmelina asiatica 
Gmelina villosa 
Gomphandra quadrifida 
var. quadrifida 
Gomphostemma curtisii 
Goniothalamus scortechinii 
Goniothalmus uvarioides 
Grewia paniculata 
Guettarda speciosa 
Gymnopetalum 
cochinchinense 
Hedyotis congesta 
Homonoia riparia 
Hopea dryobalanoides 
Horsfieldia tomentosa 
*Hoya revoluta 
Hunteria zeylanica 
Hydnocarpus castanea 
Hydnocarpus woodii 
Hydnophytum formicarum 
Hygrophila angustifolia 
Hypserpa cuspidata 
*Ilex maingayi 
Ixora brunonis 
Ixora congesta 
Ixora grandifolia 
Ixora lobbii var. 
stenophylla 
Ixora nigricans 
*Ixora scortechinii 
Jasminum bifarium 
Justicia vasculosa 
Kibara chartacea 
Knema cinerea var. 
patentinervia 
Knema cinerea var. rubens 
Knoxia corymbosa 
Koiledepas longifolium 
*Kopsia macrophylla 
Lantana camara var. 
aculeata 
Lasianthus stipularis 
Lepistemon binectariferum 
Leucaena leucocephala 
Leucas zeylanica 
*Lindera concinna 
Lithocarpus elegans 
Lithocarpus urceolaris 
Litsea polyantha 
Ludwigia hyssopifolia 
Luvunga eleutherandra 
Maesa striata 
Mallotus eriocarpus 
*Mallotus griffithianus 
Mallotus oblongifolius 
Mallotus peltatus 
Mallotus philippensis 
Mallotus repandus 


196 


Dicotyledons (cont.) 


Medinilla crassifolia var. 
hasseltii 
Melaleuca cajuputi 
Melanolepis multiglandulosa 
Melastoma polyanthum 
*Memecylon acuminatum 
Memecylon oleaefolium 
Mesua ferrea 
Millettia hemsleyana 
Millettia sericea 
*Mitrephora maingayi 
Moghania strobilifera 
Naravelia laurifolia 
Nauclea junghuhnili 
Neesia synandra 
Neonauclea calycina 
Nepenthes sp. 
Ophiorrhiza communis 
Palaquium obovatum 
Palaquium ottolanderi 
Parameria polyneura 
Parashorea lucida 
Pavetta indica 
Pavetta naucleiflora 
Peltophorum pterocarpum 
Phanera integrifolia 
Phoebe lanceolata 
Piper boehmeriaefolium 
Piper caninum 
*Piper porphyrophyllum 
Piper retrofractum 
Pittosporum ferrugineum 
Polyalthia cauliflora var. 
beccaril 
Polyalthia cinnamomea 
*Polyalthia hypogaea 
Polyalthia motleyana var. 
glabrescens 
Polyalthia rumphii 
Polyalthia stenopetala 
Polygonum chinense 
Premna pyramidata 
Psychotria rostrata 
Psychotria sarmentosa 
Psychotria viridiflora 
Pterygota alata 
Pterisanthes coriacea 
Rauvolfia perakensis 
Reissantia indica 
Rhododendron longiflorum 
var. longiflorum 
Rhynchoglossum obliquum 
Rinorea anguifera 
Rinorea macrophylla 
Sageraea elliptica 
Sauropus brevipes 
*Schefflera elegans 
Shorea guiso 
Shorea leprosula 
Shorea ovalis 
Solanum decemdentatum 
Spondias dulcis 
Stauranthera umbrosa 
Streblus laxiflorus 


Gardens’ Bulletin, Singapore — XXX (1977) 


Tamarindus indica 
*Tarenna appressa 
Tetracera scandens 
Thespesia populnea 
Thunbergia fragans var. 
javanica 
Tinomiscium petiolare 
Tinospora crispa 
Trema orientalis 
Tristania merguensis 
*Turraea brevitlora 
Urophyllum corymbosum 
Urophyllum glabrum 
Utricularia minutissima 
Uvaria cordata 
Uvaria javana 
Viburnum sambucinum 
Vitex pubescens 
Vitis lanceolaria 
Vitis martinelli 
Vitis novemfolia 
Vitis peduncularis 
Wikstroemia polyantha 
Xanthophyllum glaucum 
*Xerospermum wallichii 
Xylocarpus granatum 
Xylopia malayana 
Zippelia begoniaefolia 


MONOCOTYLEDONS 


Acampe longifolia 
*Achasma macrocheilos 
Achasma megalocheilos 
Achasma triorgyale 
Aerides odoratum 
Agrostophyllum hasseltii 
Agrostophyllum majus 
*Amomum biflorum 
*Anadendrum latifolium 
Anadendrum montanum 
Aneilema nudiflorum 
Appendicula anceps 
Appendicula cornuta 
Appendicula undulata 
Areca triandra 
Arenga pinnata 
Ascochilopsis myosurus 
Bulbophyllum pulchellum 
Bulbophyllum sessile 
*Calamus ornatus var. 
horridus 
Carex perakensis 
Coelogyne rochussenii 
*Corymborchis rhytidocarpa 
Costus globosus 
Costus speciosus 
Crinum defixum 
*Cryptocoryne affinis 
*Cryptocoryne minima 
*Cryptocoryne purpurea 
Curculigo latifolia 
Cymbidium dayanum 
Cyperus trialatus 


*Endemic to Malaya. 


Cyrtosperma lasioides 
Dendrobium acerosum 
Dendrobium excavatum 
Dendrobium farmeri 
Dendrobium indivisum 
Dendrobium leonis 
Dendrobium planibulbe 
Dendrobium pumilum 
Dendrobium secundum 
Dendrobium subulatum 
*Dendrobium tetrodon 
Dioscorea hispida 
Dioscorea polyclades 
Dioscorea prainiana 
Dioscorea pyrifolia 
Dipodium pictum 
Donax grandis 
*Dracaena porteri 
Ephemerantha fimbriata 
Epipremnopsis media 
Eria citrina 
Eria leiophylla 
Eria nutans 
Eria pannea 
Eria vestita 
Eurycles sylvestris 
Geodorum citrinum 
Globba patens 
Gymnosiphon aphyllus 
Hippeophylum scortechinii 
Homalomena griffithii 
Homalomena humilis 
Homalomena rubra 
Iguanura geonomaeformis 
Ischaemum indicum 
Ischaemum timorense 
Leptaspis urceolata 
*Licuala modesta 
Liparis compressa 
Livistona saribus 
Malaxis latifolia 
Malaxis micrantha 
Musa malaccensis 
Oberonia anceps 
Oberonia dissitiflora 
Oberonia spathulata 
Onchosperma horridum 
*Orchidantha longiflora 
Pandanus odoratissimus 
Pandanus recurvatus 
Panicum sarmentosum 
Paphiopedilum lowili 
Paspalum conjugatum 
Peliosanthes violacea 
*Phaeomeria maingayi 
Plectocomia griffithii 
Poaephyllum pauciflorum 
Podochilus tenuis 
Pollia sorzogonensis 
Pollia thyrsiflora 
Polytrias amaura 
Pomatocalpa kunstleri 
Pomatocalpa latifolium 
*Pothos latifolius 


i 


rc. .. 


Malayan Limestone Flora 197 


Monocotyledons (cont.) 


Raphidophora beccarii Setaria palmifolia Thelasis carinata 
Raphidophora korthalsii Staurochilus fasciatus Thrixspermum album 
*Raphidophora maingayi Stenotaphrum helferi Thrixspermum amplexicaulis 
Renanthera elongata Tacca leontopetaloides Th 1 : 
Sarcanthus machadonis Taeniopyllum culciferum armen pics 
Schoenorchis micrantha Taeniophyllum filiforme Trichoglottis musera 
Scindapsus scortechinii Taeniophyllum obtusum Tropidia curculigoides 
Scleria purpurascens Thecostete alata *Zingiber spectabile 


WEEDS AND ESCAPES 


PTERIDOPHYTES Erechtites Nagi tania age Solanum torvum 
. Eupatorium odoratum Spondias pinnata 
seserecoiirms Nae caaian Euphorbia hirta Tridax procumbens 
GYMNOSPERMS Flacourtia jangomas Urena lobata 
af Hyptis rhomboidea Vernonia cinerea 
Ph ag savant 
Mikania cordata MONOCOTYLEDONS 
Muntingia calabura P tetas 
DICOTYLEDONS Murraya koenigii Chrysopogon aciculatus 
Abutilon indicum ssp. Passiflora foetida var. Coix lacryma-jobi 
indicum hispida Colocasia antiquorum 
Acalypha lanceolata Physalis minima Cyperus kyllingia 
Ageratum conyzoides Pilea microphylla Digitaria violascens 
Aleurites moluccana Piper nigrum Eleusine indica 
Bidens pilosa Scoparia dulcis Imperata cylindrica var. 
Boehmeria nivea Solanum ferox major 
Cyathula prostrata Solanum nigrum Rhoeo spathacea 


TABLE 3. AFFINITIES OF THE FLORISTIC COMPONENTS 
NUMBER OF SPECIES 


ee eet a 
EXCLUSIVES sg ema 182 62 257 21.1 
PREFERENTS 9 — 53 19 81 6.6 
INDIFFERENTS 27 2 285 104 415 34.1 
STRANGERS 45 3 257 131 424 34.9 
WEEDS AND ESCAPES 1 — 29 9 39 3.2 


The species known in Malaya only from limestone (exclusives, which 
includes endemic and non-endemics) total 257. This is an increase of 62 over 
Henderson’s (1939) figure of 195. However the percentage which is 21.1% of 
the present total is less than Henderson’s figure of 26% of his total. This is 
because of the even greater increase in the overall total (from 745 to 1216) of 
the species recorded from the limestone. The total number of species endemic and 
restricted to the limestone remains unchanged with no new additions. Henderson 
recorded about 130 species, the present figure is exactly 129 species. 


The characteristic species of the limestone which are the exclusives and 
preferents number 338. This is 27.7% of the total flora. The indifferents and 
strangers are represented by 839 species (69.0% of the total flora), forming the 
bulk of the flora. The weeds and escapes are poorly represented by only 39 
species (3.2% of the total flora). 


198 


Gardens’ Bulletin, Singapore — XXX (1977) 


SECTION If — The Flora 
PTERIDOPHYTES 


INTRODUCTORY KEY TO THE FERNS 


The numbers on the right hand side refer to the numbers in the main key 
with which one should continue. 


1. Fronds of two distinct forms; nest and foliage 
1. Fronds not as above. 


. Coendveset 2. Drynaria spp. 


2. Filmy ferns; laminar part of fronds one cell thick .............,ccsssesessencees 
« sa» MRRGERTEEY ~ee ber eeed> + <3 +a.05n05 -neethathen toe: on eeeeeeee ene 6. Trichomanes spp. 


2. Not filmy ferns; fronds thicker, usually several cells thick. 


3. Fronds twining 


4. Fronds simple, erect, grass-like 


os nhawe vn DEORE OTE MEIC dg tlle pee 11. Lygodium spp. 
3. Fronds not twining. 


4. Fronds not as above. 


5. Fronds simple, entire or lobed. 


6. Fronds entire. 


ci 


ey ee yr 12. Schizaea inopinata 


Base of fronds cordate or sagittate, stipitate. ......... 16 
Athyrium cordifolium, Hemionitis arifolia, Doryop- 
teris spp. 


Base of fronds simple, if cordate then sessile. 


8. Stipe articulated 
8. Stipe if present, not articulated. 


9: 


oy 


UTIs 19. Oleandra undulata 


Sori marginal or sub-marginal .................. 
MAY osereoey or) re eee ee: 20. Vittaria spp. 


Sori not marginal or sub-marginal. 


10. 


10. 


Sori indusiate 
heterophylla, 


OTERO 23. Humata 


Asplenium spp. (some) 


Sori non-indusiate. 


‘: 
11. 


Sori? acrostichoid ......4gekea 26. 
Sori not acrostichoid. 


12. 


Z. 


Sori elongate along veins ...... 


» ea, Tene 28. Loxo- 


gramme spp., Antrophyum spp. 
Sori not elongate. 


13. 


3, 


Frond pubescent with 
stellate hairs ............ 3. 
Pyrosia spp. . 
Frond not so pubescent 
PET yy ee Eee 39. 
Lemmaphyllum_ accedens, 
Lepisorus longifolius, 
Microsorium spp. Cryp- 
sinus enervis. 


Malayan Limestone Flora 199 


6. Fronds lobed to more than half way to rachis. 
14. Sori only at the end of lobes ........................ 43. 
Ctenopteris alata 
14. Sori differently arranged. 


Ne ee Sidr tanned Source enittemaras«.: 45. 
Humata pectinata, Doryopteris spp. 
i ne 47. 


Polypodium papillosum, Phymatodes spp. 
5. Fronds compound, pinnate or more divided. 


16. Undersurface of fronds chalky-white .........0...00..0... 50. 
Cheilanthes farinosa, Pityrogramma calomelanos 

16. Not so. 
17. Sori acrostichoid or sub-acrostichoid ............ 52. 


Photinopteris speciosa, Heterogonium spp. 
17. Sori differently arranged. 
18. Sori marginal or sub-marginal, elongate ... 55. 
Lindsaya lucida, Pteridium spp., Pteris spp. 
18. Sori not marginal or sub-marginal, if marginal 
or sub-marginal, then not elongate. 
19. Sori marginal or sub-marginal. 
20. Sori on reflexed marginal flaps 
Soaked i dhcp Git plane 65. Adiantum spp. 
20. Sori otherwise. 
21. Fronds simply pinnate ...... 70. 


Ctenopteris moulton, Nephro- 
lepis spp. (some) 


21. Fronds more divided ...... 73. 
19. Sori not marginal or sub-marginal. 
aPC ios cna ci dass ot sess one suss oeak i: 


Taenitis blechnoides, Blechnum finlay- 
sonianum, Athyrium spp., Asplenium 
spp. (some) 

22. Sori rounded. 


23. Sori indusiate. 
24. Fronds pinnate or bipinnati- 
fid 86. 


Polystichum lindsaefolium, 
Pteridrys syrmatica, Cyclo- 
peltis crenata, Nephrolepis 
spp. (some), Thelypteris im- 
mersa, Cyclosorus spp. 


24. Fronds bipinnate or more 


amply divided ............ 95. 
Tectaria spp., Méicrolepia 
speluncae 

23. Sori non-indusiate ............... 101. 


Abacopteris urophylla, Acrypteris 
irregularis 


16. 


Gardens’ Bulletin, Singapore — XXX (1977) 


FERNS — MAIN KEY 


Fronds of two distinct forms, nest and foliage leaves .................esee0es 2 
Fronds not differentiated into nest and foliage leaves ....................068+ 2 
Nest leaves lobed 2 cm OF More. dlQD 5... u.sscrsc++cocvscnessanneneeennene 2 
Nest leaves not or only very shallowly scalloped ......... Drynaria bonii 
Foliage leaves pinnate; pinnae stalked .................. Drynaria rigidula 
Foliage leaves 1QDE ..... ascscnexsnae+eagsie «sets oa ees peat cs ga needane aaa 4 
Rhizome covered by short appressed scales. Sori in two irregular rows 
« sid byLsald dw vida He aia PARSE GS VRE TS » oan ela anaes Drynaria_ sparsisora 
Rhizome covered by long, semi-erect scales. Sori in two regular rows 
oo veces cetubensbet tp premelurne rutys tuleuebiiersehascauenle mee Drynaria quercifolia 


Sori enclosed by the hollow base of indusium. Sporangia on an elongate 


receptacle. Filmy feist... ...<.:.2.. conasssscshes0ncssesansieunys tans <neeen 6 
Sori different, Not ‘Mlmy. Tere”... : .7<;-s2.<cdsaseus sv pees canna peer teen 10 
Rhizome.-erect; fronds tubtéd :: scm odwkl.... Trichomanes obscurum 
Rhizome creeping; fronds borne at intervals .................ceceeeeeeeeeeees 7 
Fronds simple, orbicular to broadly ovate ......... Trichomanes motleyi 
Fronds’ pinniate'‘or rfiore dissected 30200...4/0. ATI 8 
Fronds with false .Wegus cps fits ian ox sabesintensanp > nieptenenrayhonie cine 9 
Fronds. with no. false VeqHie jes. -20sne>5 cdesceydewersnnes Trichomanes humile 
Soral-lips of two triangular lobes ............... Trichomanes bipunctatum 
Soral-lips not lobed, trumpet-like ...................65 Trichomanes christii 
Fronds. twining, Jong chim ooops sicngep sen eia ans ccweweuen pa seane tegen 11 
Fronds different: .....0..<éend uateh ee we ab WLM ees ESS Se 12 
Sterile leaflets regularly lobed; plant of forest shade .....................04 
scecseddetevabnsseeastischegaves Mian tata htaetn tee Tereeama Lygodium polystachyum 
Sterile leaflets not lobed or only the basal leaflets lobed; usually in partly 
Open places. ..... ... ice sunsnmpeeys+aseae any ae ae Lygodium flexuosum 
Fronds simple, erect, narrow and grass-like ............ Schizaea inopinata 
Fronds different... d:it-.s..rcsnderestesiowseacady est pany Ulleue ee te hein staan ttt 13 
Fronds simple, entire or lobed ............0:..<aglesdsshitas pean 14 
Fronds compound, pinnate or more divided ...............ccceceeeeeeeeeeen ones 49 
Fronds entire or almost entire, base cordate, sagittate, or simple ......... 15 
Fronds lobed, usually to more than half-way to rachis ..................06 43 © 
Frond base cordate or sagittate, stalked ........s0:...csasoessncesnntenspasvenannn 16 
Frond base not cordate or sagittate, or if appearing cordate then 
gy EEE OO oo eecepecesessosinudanaede» ty aid anne aoieine inna in 19 
Fronds: broadly: lanceolate ..............ssesesseeesreees Athrium cordifolium 


Malayan Limestone Flora 


17. 


18. 


19. 


20. 


2h. 


22. 


235 


24. 


25. 


26. 


ay. 


28. 


29. 


30. 


31. 


32. 


33. 


Stipes grooved on the adaxial side; sori superficial ... Hemionitis arifolia 
Stipes not grooved; sori sub-marginal ..................cccccceseececeeeeeeeees 


Hydathodes present as a series of sub-marginal white spots; marginal 
eNIENR IMINO fois. os. cs denn- vcs cccs cviee.oceceds Doryopteris ludens 
Hydathodes absent; marginal veins mostly jointed ... Doryopteris allenae 


Stipes articulated at a point between the rhizome and blade ............... 
eae ator eatec anata caclvaecesasrrudthacaces caste vers Oleandra undulata 
ge nn RUDD 0 FORER sey Oy Eee Poe ee Ae eae 
ee ee INNS = 000 RERIS-MUAO OTN So ss oleic ceva ig os gals inside se seese’scaupaieee 
SOF! MOC MiAPial OF SUD-NATPINAL ooo... 5 <5. cece kc cece sect eesesenceceeses 
Fronds sessile, 20 by 0.2—0.3 cm ..................05. Vittaria angustifolia. 
Fronds stipitate, to 90 by 0.5 cm ...... Vittaria elongata var angustifolia 
Stl WIGUSIOES 5s ess oce- Tevvexed: -heue- dopa vedpeeses 6 MEARE. Gal bkns -eep ane. de... 
NG Se ESE FEIT OPEL DST PDO ig at SAE 
Rhizome long creeping; fronds dimorphic ......... Humata heterophylla 
Rhizome short, erect; fronds not dimorphic .....................cceeceeeeees 
os OU. | a eee eres Cm Met i), ere Aes Oe 
Feomidis: pinnate... dizhin.ods..to. akin acidiie.ne. os Asplenium salignum 
Veins of fronds united in a sub-marginal vein ...... Asplenium phyllitidis 
Veins of frosd-all free si). 225). becca). cbes 238 Asplenium squamulatum 
Sori acrostichoid or sub-acrostichoid ............... Leptochilus decurrens 
Sori not acrostichoid or sub-acrostichoid ..................ecececeeeceeeeeceees 
Suen ere tS MCE ONC CL ST.  SUL Sater cacne oun wclc os indcs-cuae.cecccasese 
Sent Mot elomeet- lone: Veins! 20.22. eceicc ree Pe 
Sori obique to the midrib, in roughly parallel rows ......................4. 
Sori along many veins, forming a network ......................ccceeeeeeeeees 
Midrib raised on the lower surface, almost flat above ..................... 
STE AE, / OLE EEE DENS EE Pe Ee Loxogramme scolopendrina 
Midrib raised on the upper surface, almost flat below .....................55 
aN eo ius tl ends beh each atdbenttalindees os oe Loxogramme avenia 
Parapevees Mi SOT Club-shaped  .. +... ... 0. feauccars asp - Wide ory bees omar ones 
Paraphyses in sori thread-like ..................... Antrophyum callifolium 
Stipes to 10 cm; fronds to 30 by 10 cm...... Antrophyum semicostatum 
Stipes not distinct; fronds smaller, to 15 by 2 cm ....................020ee 
ergy Gap map lee os Reka) sageny age 7k See ae Antrophyum  parvulum 


Fronds densely or sparsely covered by stellate hairs, more on the lower 
NUNC a hciees - eye < Lge atete Chbice © Beg. nae ees - 45+ cage paeravae ablaseli gees pie eee 


eee RS SCT POMPE Cla) DTOAG oo. i. li ccc ducectcccteccccecccetececess 
Sterile fronds orbicular to broadly ovate ...... Pyrrosia nummularifolia 


201 


18 


20 


21 
22 


23 
26 


24 
25 


27 


28 
32 


29 
30 


31 


33 
39 


34 


202 


34. 


35. 


36. 


ay; 


38. 


39. 


40. 


41. 


42. 


43. 


45. 


46. 


Gardens’ Bulletin, Singapore — XXX (1977) 


Sori in a single row on either side of the midrib, to about 0.5 cm across 
spediepeepipesssoreagcsaese regs snennnnceeienpeareaesh eee Pyrrosia angustata 


Fronds dimorphic; sterile shorter and sometimes wider ..................- 
Fronds not dimorphic; sterile about the same size and shape ............ 


Sterile fronds 1 — 1.5 cm wide; fronds when dried, the margins involuted 
the upper surface becoming concave; scales of rhizome not closely 
BPPTESSEU soo... ; ssa ncilelhegoneceeeasepeet saan aen eee Pyrrosia adnascens 
Sterile fronds 2 — 3.5 cm wide; fronds sine dried, the margins revoluted, 
the upper surface becoming convex; scales of rhizome not closely 
Spprested weiss Liv. tohe.. Heard Tia bods. cet kee Pyrrosia varia 
Lower surface of frond densely brown-stellate pubescent .................. 
a 8's aewad dunainw ss seine getline Pachbanviag .eaiaal he tee cane ntcere: Manian Pyrrosia stigmosa 


Fronds about 1.8 cm wide; sori on the apical half or whole of frond; no 
clear marginat the. edge... 200/42. 228OT.. Sa Pyrrosia floccigera 
Fronds 3-6 cm wide; sori on the apical third of frond; with a clear 
margin to 1 cm wide at the edge ..................... Pyrrosia penangiana 
Sori in two rows, one on either side of the midrib ........................04. 
Sori scattered all over the surface ...............cceceeeeeees peat - 2p ope ends 
Fertile part of frond narrowed; frond less than 15 cm long ............... 
eT ee ee Te mes ee seeseeeee. Lemmaphyllum accedens 
Fertile part of frond not narrowed; frond to over 30 cm long ............... 
a veeeas Sande wqdqaigomninee tepiliaey atte aaate net ren Ramee meee ... Lepisorus longifolius 
Fronds stipitate’:. i5..52.0c2.5000<s9) tesa. cs appeee aatOle ene ee 
Fronds sessile, widening abruptly from the base: .s5..:ipncis ie ee 
winie ankceo dadlee og tes eg gree eaenio a en aeececeseeccsseeeeeeeese, Microsorium musifolium 


Stipes 1 cm or less; fronds widening very gradually from the base ......... 


svevasecvedddeuin elle deed DEW adelc AED Le hDGIE ENE iceEmanne Microsorium punctatum — 


Stipes 5 cm or more; fronds with broad bases ......... Crypsinus enervis 


Sori only at the ends of the lobes of frond; usually 2—5 on each lobe 
oy wo sibs «win dialuidcaspinaty 5 #G'anitidinenela peal P Ena Ctenopteris alata 


Sori continuous along margins of fronds .................csceeeceeeeceeneneeees 
Sori not so; in two rows, one on either side of midrib .....................06. 


Hydathodes present as a series of submarginal white spots; marginal 
veins mostly free. o.4).0:30000 BAER. AR, RE Doryopteris ludens 
Hydathodes not present; marginal veins mostly joined ................+++ 
a ct nacadbescuscescp scenes epiinscnsnds wees ashes sitet 


35 


36 
37 


38 


40 
41 


42 


47 
46 


Malayan Limestone Flora 


47. 


48. 


49. 


50. 


51. 


52. 


a: 


54. 


Do- 


56. 


57. 


58. 


=. B 


Veins anastomosing. Rhizome stout, to over 1 cm diameter ............... 
Veins all free, forked once. Rhizome slender, 0.2 —- 0.3 cm diameter ...... 
Se epee Htabebrets OES we seta den cng 500 asesseeeeeee Polypodium papillosum 


Fronds with 1-4 pairs of lobes; sometimes simple ........................ 
0 he le Baie ate eee ps ae Teo et fe a Phymatodes_ scolopendria 
Fronds with 7-10 pairs of lobes; never simple .......................:.:0ee 
ensaeaene teases 2s als ass ss, ha 2k4 css ia snd ves Phymatodes nigrescens 


Underside of fronds completely or partially covered by chalky-white 


SON Metta ies canna te celduiea Laka SCAM IEUAGLOY. 2352. L0se 220.00. Wk aa. 
Underside of fronds not chalky-white ...................ccccceeeeceeeeeeeeeeees 
Fronds to 20 cm long, often less; sporangia near the margin of frond 
en 5 Ra eee ee Cheilanthes farinosa 
Fronds much larger, 30-60 cm long; sporangia scattered all over the 
eeeeTertMCe OF LOU oo. 5.0.0 cea> os 4 ye3ca8e5 «* Pityrogramma calomelanos 
Sori acrostichoid or sub-acrostichoid, on the apical part of frond or 
IRIN: OSM: SCN isi ea ae ria dhs inset Sun somes dignasicanenedss- 
Sori not acrostichoid or sub-acrostichoid ..................ccceececeeeeeeee ces 
ra OMT A AWG A RR ON. Photinopteris speciosa 
ED uss <4 sccawans sx onbue anew hide dass wied ove kes. dUees (NEL ee TOI 
Srcemteems ct SAY Ce) IO Heterogonium alderwereltii 
oR aa ee eC Se ne Heterogonium pinnatum 
Sori marginal or submarginal, continuous in a short or long band ...... 


Sori not marginal or submarginal, if marginal or submarginal, rounded 
I CN SE Sean Seat ace teak twink suas anus hcthckoaensasesesesscese 


Sori short banded, uniting the end of several veins ...... Lindsaya lucida 
Seed > AUN Swans ik sannenapainas RN Mette adh inns ns hgndy <5 
Fronds tripinnate or quadripinnatifid; of long continued growth ......... 


Fronds pinnate or tripinnatifid or rarely tripartite; not of long continuous 
cla rig ce a ect ga le nea pg SRSA UOT ~ SELEY ICA ask AA 0 


Ultimate segments widely spaced, connected by a wing decurrent from 


the preceeding segment ............ Pteridium caudatum var. yarrabense 
Ultimate segments close together, not connected by a wing ............... 
Ye, BUTE aeeesescseeeseeesscessseees Pleridium aquilinum var. wightianum 
PMUM RRR MNII, OUMEAIN S556) Syd ca cdccaasescscccescscsvescess Pteris vittata 
a aN coats oc ock ss cdlacccqcccbdcccsscwandedqedsasseeesse 


Fronds dimorphic in mode of branching; sterile, with deeply lobed 
pinnae; fertile, with only the basal pinnae having a single basiscopic 
IR Fe Od. as svat bans deséadsceadovewe Pteris ensiformis 


50 
51 


52 
54 


53 


55 


63 


56 
57 


58 


59 


204 


60. 


61. 


62. 


63. 


64. 


65. 


66. 


67. 


68. 


69. 


70. 


7h. 


TZ, 


te 


74. 


7. 


Gardens’ Bulletin, Singapore — XXX (1977) 


Pinnae, margin entire or serrulate .................eceeeeees Pteris scabripes 
Pinnae;: deeply: lobed ©. .2aeasle. ausessichs | «an 065% Seon ae bo aeeN aed ene 
Fronds tripartite, basal pinnae almost as large as the terminal one ......... 
a sio-n ssln'o s,s.» 6's un o's ODAC APEEDS > Mp RE ep eetoae seit tani Oe lk een Pteris tripartita 
Fronds not tripartite. .00y.5liici. 0.200. .cetbe deel en calves dallas Danae 
Fronds less than 50 cm long; lowest vein in each pinnae lobe 3-4 times 
FORCE)... FRAO Rah A ii Re Pteris mertensioides 
Sori marginal or:submarginah: ov. . 21). .//Waviiiwa oven. 1a. Gee 
Sori not. marginal. or. submargimal ............casccsccaccesanessennsenss SRE 
Sori on small ‘reflexed margmal flaps 17.00.52... 2201.22 ee 
Sofi 1.otherwise 913. isa. sien gana. cewel- ceties «eee eee a ee 
Fronds simply pinnate «....62..2cccesccesecssees « coevrdaesieud taco teaeueenaann 
Fronds tripinnate’or more ‘divided '):.//)2..VGs......19018). SAUL, BO 
Stipes, rachises and stalks of pinnae winged ...... Adiantum soboliferum 
Stipes, rachises and stalks of pinnae not winged .................cceeeeeeeeees 
Rachis and pinnae hairy; indusial flaps almost circular ..................... 
PROT NRE OR MVOSEY ee Ger BY meg nt. hme canto ... Adiantum malesianum 
Rachis hairy on the upper surface only; pinnae glabrous; indusial flaps 
broader, than loge «iscsi vee eben huis ge easacecnees Adiantum zollingeri 
Fronds tripinnate, basal angle of leaflet usually more than a right-angle 
Joana is: -yetan aan Sts aa con poet ek ein as Adiantum stenochlamys 
Fronds quadripinnate or more divided, basal angle of leaflet usually 
less than "9. Tight-Aneye | o.0\s ccwecencs oneav aden beta meaea et Adiantum tenerum 
Fronds sunply -qeniiate.:.3.9.5.:4-a3ue anes Te ey Rr 
Fronds tripinnatifid..or more :dswided) sic. ..cs. aegis idan se seeemanaeiee ane 
Sori indusiate; fronds 30 Ci OF MORE Jecschevaccevtes beige .cdsee cr Pearene ieee 
Sori non-indusiate; fronds ;10-20 sont oo) Aaisinsfeesmendcnen Doane eee 
snap sun vhele cece s Deglsall seve yiteee Ctenopteris moultoni, Nephrolepis radicans 
Apex of ‘pinnae ‘roundéd/7), Sarat na aan Nephrolepis radicans 
Apex. of. pinnae. pointed: .ii.:<iaianes oaeeke y= dbanenbbite tonnciieasan eens 
Pinnae about 5 cm long, strongly falcate ............... Nephrolepis falcata 
Pinnae about 8 cm long, weakly falcate ............ Nephrolepis hirsutula 
Fronds tripinnatifid or tripinnate, no false veins; sori longer than broad 
sbidal aphes .  Jalvs SoU ey ae Wes ecessvererseedecverepess aoe 
Fronds quadripinnate or more divided, false veins present; sori about as 
Wide as J6mg 4,5. Rhos: eee eee Davallia denticulata 
port clompate « . j.i:nincmiv eee ean 1-5 colws'to ees o'vislers wecalniabnelen «heen 
Sori ‘rounded!’ /,, 211212, .20j0a0ek Le sbleinn shoetGleodhk. AUST IOILU: AenGnnne! 
Sori in a longitudinal band halfway between the margin and costa ...... 
saEON eee gE Sun bie he ve »3'y 05.3 ve'cbG bobs COMER AMAA POREr ee ae Taenitis blechnoides 
Sori different | »........cscsceeredseietedesscueeeaddvedsvoth adelante 


61 


62 


64 
74 


65 
69 


66 
68 


67 


70 
73 


71 


Te 


75 
84 


Malayan Limestone Flora 


76. 


fee 


78. 


79. 


80. 


81. 


82. 


83. 


84. 


85. 


86. 


87. 


88. 


89. 


90. 


Sori elongate along and on either side of the midrib ........................... 
We0o Ait. Ot. bnidieiwt. lon endeal cv. t2cu. Blechnum _finlaysonianum 


Sori, on one or both sides of vein; transverse section of the upper part 
of stipe shows vascular strand with two arms (easily seen in fresh 
ak a ead ia en ert ce, SER obs ce Ed « cigs anas benes@dsves vee petees 
Sori, all on one side of vein; transverse section of the upper part of stipe 


aiowe vascular strand: with! four AMIS: -.20i. 52s. k ii... oo. ce ec ceceeccu ens 
Fronds simply pmmnate; pinnae entire or lobed ....................s..cccseseces 
Fronds more amply divided. Large plant, stipe and frond to 2 m long 
LEC SLI SP Pa OF Se aCe ey eee asceeeeseseeeee. Alhyrium esculentum 
a, Oe a ae es Se 
RS BIER TOQUE a ick. wiry Degen iy Scenes oh hyans xe Athyrium pinnatum 


Lateral veins of pinnae forked once, basiscopic veinlet simple or forked 


a SNE a OE AS a oe a OL Athyrium montanum 
Lateral veins of pinnae in clinic groups, each with 3-4 pairs of veinlets 
(LA Snel eg lenge gamain, Hope be opt Ba ede Athyrium prescottianum 
Rhizome slender, creeping; fronds at interval ...... Asplenium unilaterale 
Rhizome stout, short creeping; fronds tufted ......................0..cce cee 
Lower pinnae gradually reduced .................. Asplenium pellucidum 
Lower pinnae not reduced ..................... AD CAINS. SOA... 
PaO OY 2 CID sacainw stn ndanpectne nyt Pub nh Rg Asplenium adiantoides 
ie OA OY GHD oon i acce cen nsh acne pees Asplenium macrophyllum 
Sori indusiate; indusia sometimes deciduous ................ccccccccccceceeceeee 
Son non-iIndusiate 2... 250.7 00.00. eo a a bs Re, | he a9 ae Ee ie 
Peonds giniate Or at most bipiniatifid © ............. 0.05.0. 000. 0c. cece cceceees 
Fronds bipinnate or more amply divided ................... 0... cccceceeeeeeee es 
EOGES SO 2D CIN WHEE oo 5. dain dvi ges los nae0’ Polystichum lindsaeifolium 
an tt A WN HCN AS . CAPS ss cis acs cia de Gdaccadecaceac 
A tooth is present in the sinus between lobes ......... Pteridrys syrmatica 
a ea cue sdglegewaumené 
Sori in three irregular rows on either side of the midrib .................. 
i Easy een Baty oy oO Caen APY Reon, OP Ce Cyclopeltis crenata 
I MLR scence ndvcavacncessscecucscnsutcccccasteres 
Ot e@ftaneed On elmer side, Of the. cOSsta.. .......... 00. <rasdececmaneemaracesiles. 
Ot atraticed On,either Side of the costule ..............cccccccccascsecccseccs 
Sor ‘on- marginal lobes sii! 30. .tv. eee. Nephrolepis dicksonioides 


Sori not on marginal lobes ......................0000 Nephrolepis biserrata 


205 


77 


78 


8] 
79 


80 


82 


83 


85 
101 


86 


95 


87 


88 


89 


90 
91 


206 


91. 


92. 


oa: 


94. 


95. 


96. 


O7. 


98. 


99. 


100. 


101. 


Gardens’ Bulletin, Singapore — XXX (1977) 


Veins of the adjacent lobes of pinnae fused to form an excurrent vein in 
the sinus. If this is not distinct then lobes only two-third to the costae 
vapuneeb vd ead actacdionscweubbabeesilen Oh dhWivea Ne O4E POM MOONE cUgth ERI An ian ti inn 92 
Veins of the adjacent lobes of pinnae not fused, at the most meeting at 
the sinus. Lobes almost to the costae ............... Thelypteris immersa 


Sori confined to the lobes of the pinnae, not on the lower veins ......... 93 
Sori not confined to the lobes of the pinnae, also on the lower veins ...... 94 


Pinnae mostly lobed two-third to the costa. Lowest pair of pinnae not 
TOUUICG se ravcauees <0 avienasn oats foo tee eae eee ee Cyclosorus extensus 
Pinnae lobed one-quarter to one-third to the costa. Lowest pair of pinnae 
often Teducea tay auricles 70 eee Cyclosorus interruptus 


Pinnae to 1 cm wide; costae densely pubescent beneath ..................... 
aa scnaceanainomaye 4 gn'dainy Wha ae Sg pein Say Ueto Mla ea Cyclosorus unitus 
Pinnae to 3 cm wide; costae minutely pubescent below ..................... 
catresentsWactstesth goss ieee teen tt eo tee an ame aan name Cyclosorus megaphyllus 


Basiscopic lobe of the lowest pair of pinnae enlarged .....................04. 96 
Not so. Fronds deeply dissected: $0.0 2005. es .cs. «ses ape atadan eee een 100 


All sori on anastomosing VeINS .................0ceee eee Tectaria amplifolia 
All or nearly all sori. on.free ‘yeims i460: dens sndlespimsenes heres ea 97 


Fronds glabrous on the upper surface, or with some hairs near the 
SUMUSES 52, - o mel'e cae g guidisen + ac te eo atonbecun ly snure bald wate c/a age ta a 98 


shee aga sie aieladin ae de Dee otaama i aan ane Miaka a Tectaria_ variolosa 
Scales stiff, dark purple-brown. Fertile frond not contracted ............... 
gh nu o-nee ¢ouegecdll oe ¢ o/c ohielyOil te tins tain aetnane nanan Tectaria griffithii 


Veins anastomosing in the costal and costular areoles only (forming a 
single row of areoles on either side of the costae and costules) ............ 
saia's sian apn! wid cle tall» Aa Sanaa ky Ae aida te assesses. Tectaria devexa 


Under-surface of lamina glabrous, glabrecent or rarely pubescent ......... 
OTETOTTT TPE CA hy Tr FEY Pe ei Microlepia_ speluncae 
Under-surface of lamina densely pubescent .................c.sceceeeeeeeee ones 
ea le ciated se ne Microlepia speluncae var. villosissima 


Pinnae not lobed, at most crenate or toothed only ...............cececeeeeees 
5 Pees Wide bon 0 Zsiaiv'c b's snes Sn gale loyal ea wees. Abacopteris urophylla 
Pinnae deeply lobed, often pinnate at the base; basiscopic lobe of lowest 
Pinnne -Calarged 3.0)... .45..ssiasdessesesserduesesselieene Arcypteris irregularis 


Malayan Limestone Flora 207 


Adiantaceae 


Adiantum malesianum Ghatak, Bull. Bot. Surv. Ind. 5: 73. 1963; Holtt., Fl. Mal. 
2: 638. 1966. 


A. caudatum L., Mant. (1771) 308; Holtt., lc. 599. 
Distributed in N.E. India, S. China, Thailand, Indo-China, Sumatra, Sarawak 


and Philippines. Common on limestone in Malaya but also found on other rock 
types. A fern of moderately shaded places. 


Adiantum soboliferum Wall. apud Hk., Spec. Fil. 2: 13. 1851; Holtt., Fl. Mal. 
2: 598. 1966. 


Distributed throughout the Old World tropics. In Malaya recorded only from 
the north and usually on limestone. 
Adiantum stenochalamys Bak., Ann Bot. 5 : 29. 1891; Holtt., Fl. Mal. 2 : 602. 1966. 
A. opacum Copel., Phil. J. Sc. 1, Suppl. 255, t. 3. 1906. 


In Malaya, nearly always found on rocks near the sea. Found inland only on 
rocks along the Tahan river and on limestone along Sungei Betis in Kelantan. 


Adiantum tenerum Sw., in Hk. et Bk., Syn. Fil. (1868) 124; v.A.v.R., Mal. Ferns 
(1908) 330. 
A tropical American species now widely cultivated in tropical, subtropical or 


even temperate countries. Recorded on limestone as escapes from cultivation; on 
shady hill slopes. 


Adiantum zollingeri Mett. ex Kuhn, Ann. Mus. Bot. Lugd. Bat. 4: 280. 1869; 
Holtt., Fl. Mal. 2: 638. 1966. 


A. caudatum L. var. subglabrum Holtt., l.c. 600. 


Distributed in Ceylon, S. India, Thailand and Indochina. In Malaya recorded 
only from limestone in Kedah and Perlis. 


Antrophyum callifolium Bl., Enum. Pl. Jav. (1828) 111; Holtt., Fl. Mal. 2: 605. 
1966. 


A common forest fern, epiphytic or on rocks. Recorded from limestone in 
South Kelantan and Central Pahang. 


Antrophyum parvulum BI]. Enum. PI. Jav. (1828) 110; Holtt., Fl. Mal. 2 : 605. 1966. 

Distributed in Java. In Malaya this species is widely distributed and common 
on limestone, on rocks; with only one record as an epiphyte (Henderson 1939). 
It has been recorded from limestone in the south in Johore and apparently absent 
from the vast tract of forest separating the Johore from the northern limestone. 
It has been collected away from limestone only twice, both on Penang Hill. 


Antrophyum semicostatum Bl., Enum. Pl. Jav. (1828) 110; Holtt., Fl. Mal. 2: 605. 
1966. 


An epiphytic fern of mountain forest. Recorded from low elevation on lime- 
stone probably as epiphytes in Kelantan. Resembles A. callifolium but distinguished 
by the club-shaped paraphyses. 


Cheilanthes farinosa (Forsk.) KLf., Enum. Fil. (1824) 202; Holtt., Fl. Mal. 2 : 592. 
1966. 
Pteris farinosa Frosk., Fl. Aegypt. Arab. (1775) 187. 
Aleuritopteris farinosa Fée, Gen. Fil. (1850-52) 153. 


208 Gardens’ Bulletin, Singapore — XXX (1977) 


Distributed in most tropical and temperate regions with two records from 
Malaya, both from limestone. One was from Bukit Baling in Kedah and the other 
from Bukit Chintamani in Pahang the Chintamani specimen was growing in rock 
crevices from vertical cliffs. 


Doryopteris ludens (Wall.) J. Sm., Hist. Fil. (1875) 289; Holtt., Fl. Mal. 2: 594. 
1966. 


Pteris ludens Wall. apud Hk., Spec. Fil. 2: 210. 1858. 


Distributed from N. India and S. China southwards through Malesia. Restrict- 
ed to limestone in Malaya and all collections except two are from Kedah and 
Perlis; the exceptions are from Gua Batu, Selangor and Batu Kurau, Perak. 


Doryopteris allenae Tryon, Contr. Gray Herb. 91:91, 97. 1962; Holtt., Fl. Mal. 
2: 638. 1966. 


This species is similar to D. ludens in appearance. It differs however in having 
the rhizome short creeping rather than long creeping, the marginal veins in the 
sterile lamina mostly jointed rather than free and in not having any hydathodes 
on the upper surface of the fronds. 


Endemic to limestone in Malaya, recorded from Perak and Selangor. Usually 
in shaded localities from rock crevices. 


Hemionitis arifolia (Burm.) Moore, Ind. Fil. (1859) 114; Holtt., Fl. Mal. 2 : 596. 
1966. 


Asplenium arifolium Burm., Fl. Ind. (1768) 213. 


Distributed in Ceylon, India, Burma, Indochina, Malesia and the Philippines. 
In Malaya only from the north, nearly always from limestone. 


Pityrogramma calomelanos (L.) Link, Handb. Gew. 3 : 20. 1833; Holtt., Fl. Mal. 
2: 593. 1966. 


Acrostichum calomelanos Linn., Sp. Pl. (1753) 1072. 


Pantropic, originating in tropical America. Common in Malaya, from the low- 
lands to 1,300 m. Frequently one of the early colonisers of open ground. Recorded 
once from the base of limestone. 


Taenitis blechnoides (Willd.) Sw., Syn. Fil. (1806) 24, 220; Bedd., Handb. 410, t. 
242; Holtt., Fl. Mal. 2: 586. 1966. 


Pteris blechnoides Willd., Phytogr. (1794) 13. 


Vittaria angustifolia Bl., Ennum. Pl. Jav. (1828) 199; Holtt., Fl. Mal. 2: 610. 1966. 


Distributed throughout Malesia. In Malaya, Sumatra, Borneo and Java this 
is a mountain epiphyte at 600-1800 m. On limestone in Malaya this has been 
collected as a low epiphyte from 20-300 m; recorded from Kedah, Kelantan and 
Perlis; uncommon. 


Vittaria elongata Sw., Syn. Fil. (1806) 302; Holtt., Fl. Mal. 2: 614, 1966. var. 
angustifolia Holtt., in l.c. 


This variety is endemic to Malaya and found only in the North-west and on 
Pulau Tioman in the South-east. It is both an epiphyte and rock plant and 
recorded from only one limestone locality as an epiphyte. The typical variety is a 
common lowland and mountain epiphyte with a paleotropical distribution. 


Malayan Limestone Flora 209 


Dennstaedtiaceae 


Arcypteris irregularis (Pr.) Holtt., Reinw. 1: 193. 1951; Fl. Mal. 2: 538. 1966. 
Polypodium irregulare Pr., Rel. Haenk. 1 : 25. 1825. 


A common forest species in Malaya. Recorded once from the small limestone 
outcrop in deep forest in Johore. 


Asplenium adiantoides (L.) C. Chr., Ind. Fil. 1905; Holtt., Fl. Mal. 2 : 431. 1966. 
Trichomanes adiantoides L., Sp. Pl. (1753) 1098. 
Distributed in Burma, Thailand and southwards to Malesia, Australia and 
Polynesia. Also in Madagascar. 


Resembles A. macrophyllum Sw. but generally smaller. Intermediates occur 
but the extreme forms are very different. More field work and cultivation of this 
species will help to clarify the growth form of this species. Presently they are best 
kept apart. 


This is a fern of rocky places and nearly always from limestone. It is very 
common on scrubby summits of some hills around Gua Musang in Kelantan. 


Asplenium macrophyllum Sw., Schrad. Journ. 1800/2: 52. 1801; Holtt., Fl. Mal. 
2: 431. 1966. 


Found in most parts of Malaya on rocks and as epiphytes. It is common on 
coastal areas in the east coast and on limestone in Selangor. 


Asplenium pellucidum Lam., Encyl. 2 : 305. 1786; Holtt., Fl. Mal. 2 : 428. 1966. 


Common in Malaya from the lowlands to the hills, on rocks or as epiphytes: 
often on limestone. 


Asplenium phyllitidis Don, Prodr. Fl. Nep. (1825) 7; Holtt., Fl. Mal. 2 : 420. 1966. 
Common in Malaya, usually epiphytic. Recorded from limestone in Kelantan. 

Asplenium salignum Bl., Enum. Pl. Jav. (1828) 175; Holtt., Fl. Mal. 2 : 421. 1966. 
A. filiceps Copel., Philip. J. Sc. Sc : 285. 1910. 


Common in Malaya, usually as an epiphyte, on the lowands, hills and moun- 
tains. Fairly common on limestone under partial shade as a low epiphyte or on 
rocks. 


Asplenium squamulatum BI., Enum. Pl. Jav. (1828) 174; Holtt., Fl. Mal. 2 : 426. 
1966. 


Distributed throughout Malesia. Recorded in Malaya from Pahang, Kelantan 
and Selangor, usually on limestone. 


Asplenium unilaterale Lam., Encys. 2 : 305. 1786; Holtt., Fl. Mal. 2 : 438. 1966. 
Widely distributed in Malaya, on rocks in moist shady places. Uncommon on 
limestone. 
Athyrium cordifolium (Bl.) Copel., Philip. J. Sc. 3c: 300. 1908; Holtt., Fl. Mal. 
2: 548. 1966. 
Diplazium cordifolium Bl., Enum. Pl. Jav. (1828) 190. 
Common in lowland and mountain forest in Malaya. Recorded only once from 
limestone. 
Athyrium esculentum (Retz.) Copel., Philip. J. Sc. 3c : 295. 1908; Holtt., Fl, Mal. 
2: 562. 1966. 
Hemionitis esculenta Retz., Obs. Bot. (1791) 38. 


Common in wet places in the lowlands of Malaya. Recorded once from lime- 
stone at the base of hill. 3 


210 Gardens’ Bulletin, Singapore — XXX (1977) 


Athyrium montanum (v.A.v.R.) Holtt., Fl. Mal. 2: 555. 1966. 
Diplazium montanum v.A.V.R., Bull. Jard. Bot. Ser. If, 28: 19. 1918. 
Common in Malaya in lowland and hill forest. Recorded from limestone in 
Johore only. 
Athyrium pinnatum (Blanco) Copel., Philip. J. Sc. 3c : 297. 1908; Holtt., Fl. Mal. 
2: 560. 1966. 
Allantodia pinnata Blanco, Fl. Filip. Ed. 2 (1845) 571. 
Widespread in Malaya with one record from limestone. 


Athyrium prescottianum (Wall.) Holtt., Fl. Mal. 2: 557. 1966. 
Asplenium prescottianum Wall., Cat. (1829) 235 (nom. nud.) 
Endemic, an uncommon species. Recorded once as an epiphyte from limestone. 


Blechnum finlaysonianum Hk. et Grev., Ic. Fil. (1831) t. 225; Holtt., Fl. Mal. 
2: 445. 1966. 
A common species in lowland and hill forest, once recorded from limestone. 


Cyclopeltis crenata (Fée) C. Chr., Ind. Fil. Suppl. 3 : 64. 1934; Holtt., Fl. Mal. 
233527, 1966, 
Hemicardion crenatum Fée, Gen. Fil. (1852) 283, t. 22A. 


Distributed in Burma, S. China and western Malesia. This is a plant of rocky 
places, usually but not always from limestone, in Malaya. Widely collected from 
limestone, in part shade. 


Davallia denticulata (Burm.) Mett., in Kuhn., Fil. Deck. (1867) 27; Holtt., Fl. Mal. 
22309. 1966. 
A common rock plant and epiphyte; recorded a number of times from lime- 
stone. 


Davallia solida (Frost.) Sw., Schrad. Journ. 1800/2: 87. 1801; Holtt., Fl. Mal. 
2: 360. 1966. 


Trichomanes solidum Forst., Prodr. (1786) 86. 


A common coastal epiphyte and rock plant in Malaya. Uncommon inland. 
Recorded only once on limestone; this was from a large local population on the 
summit of Batu Tapah, growing in part shade on rocks and as a low epiphyte. It 
is probably not uncommon on limestone, on such dry, rocky, scrubby summits. 


Heterogonium alderwereltii Holtt., Sarawak Mus. J. 5 : 163. 1949, Fl. Mal. 2 : 522. 
1966. 


Pleocnemia membranifolia p.p. quoad Bedd., Handb. Suppl. (1892) 48. 


Plant like that of H. pinnatum. The typical specimen however has larger 
sterile fronds with 4~7 pairs of pinnae. The sori is not acrostichoid but is either 
distinct or elongate along veins. The indusia is distinct though less prominent in 
mature sori. 

Distributed in Sumatra. Restricted to limestone in Malaya (except for a 
specimen from Patani, Kedah); not as common as H. pinnatum. 

For notes see under H. pinnatum. 


Heterogonium pinnatum (Copel.) Holtt., Sarawak. Mus. J. 5: 163. 1949, Fl. Mal. | 
2: 524. 1966. 
Stenosemia pinnata Copel., Phil. J. Sc. 1, Suppl. 2 : 48. 1892. 
Pleocnemia membranifolia p.p. quoad Bedd. Handb. Suppl. (1892) 48. 


___ Distributed in Sumatra, Borneo and the Philippines. Restricted to limestone 
in Malaya, and the commonest fern on limestone. 


Malayan Limestone Flora 211 


Typically this fern is distinguished from H. alderwereltii Holtt. mainly by the 
absence of indusia and the sterle fronds having only one instead of four to seven 
pairs of free lateral pinnae. H. alderwereltii has distinct sori or extended sori along 
the veins while typical H. pinnatum has acrostichoid sori, and if atypical, the sori 
extend along veins but still without indusia. 


However, there is great variation in the morphology of these two species, thus 
delimited. There are H. pinnatum plants with up to 7 pairs of free pinnae (in the 
sterile fronds) and which have, in the fertile fronds, an acrostichoid condition in 
which the sori are non-indusiate. The sterile lamina with up to 7 pairs of free 
pinnae would seem to belong to H. alderwereltii while the acrostichoid non- 
indusiate sori would seem to belong to H. pinnatum. Apart from the extreme forms 
of H. pinnatum in which the sterile fronds have normally one pair of free pinnae, 
the form and size of the sterile fronds are insufficient characters to separate these 
two species. 


There are identical plants in which the sori are distinct or elongate sub- 
marginally. In some instances they are indusiate and in others they are not. Once 
these two species are kept distinct, one can say that the non-indusiate plants would 
be abnormal H. pinnatum (the normal plants have acrostichoid sori). However, 
this argument would be equally valid if one says that the non-indusiate condition 
is found on abnormal H. alderwereltii which have lost their indusia. In both 
indusiate and non-indusiate forms there often exists a condition in which sori are 
found on fronds with the size and form of a sterile frond. These sori are either 
very distinct on vein endings or often form submarginal bands. 


This range of variation can be found in a single population such as that on the 
base of Bukit Takun, Selangor, growing on limestone and organic debris. Only 
the extreme forms would seem to keep these two species apart and I suspect (as 
Beddome thought) they are variations of one species. This is as much as I can 
conclude from field observations; experimental culture from spores is required. 


Besides these variations there are indusiate as well as non-indusiate forms 
which are free veined, a character now found in H. sagenoides (Mett.) Holtt. Also, 
the form in which the fertile fronds have a broad lamina and distinct sori 
encroaches on one of the characters of the remaining species of Heterogonium 
found in Malaya, H. saxicolum (Bl.) Holtt. Other characters, however, seem to 
keep H. pinnatum and H. alderwereltii distinct from these latter two species. The 
former two are also different in being exclusively limestone plants in Malaya with 
an exception of one specimen of H. alderwereltii collected from Patani, Kedah. 


In the light of recent collections, the taxonomic status of this complex of 
Heterogonium needs looking into, especially the limestone-inhabiting groups. 


For convenience, the two species are retained here, often distinguished only 
by the presence or absence of an indusium. H. pinnatum is the more commonly 
collected species, appearing on most limestone outcrops except in the extreme 
north and Kelantan. Both are found in shady, rocky places and most luxuriently 
in situations with humus accumulation. 


It is interesting to note that H. pinnatum has been collected from the very small 
and isolated Johore limestone, forming a link with the more southern Sarawak 
populations. 


Humata heterophylla (Sm.) Desv., Prodr. (1825) 323; Holtt., Fl. Mal. 2 : 366. 1966. 
Davillia heterophylla Sm., Mem. Ac. Turin. 5: 415. 1793. 


A widely distributed species in Malaya usually in exposed places, on rocks 
or as epiphytes. Recorded on limestone from Kelantan. 


212 Gardens’ Bulletin, Singapore — XXX (1977) 


Humata pectinata (Sm.) Desv., Prodr. (1827) 323; Alston, Phil. J. Sc. 50: 175 
1933; Holtt., Fl. Mal. 2 : 369. 1966. 


Davallia pectinata Sm., Mem. Ac. Turin. 5 : 415. 1793. 
Distributed from Sumatra to New Guinea, usually on coastal rocks and trees. 
Inland collections in Malaya are restricted to limestone, growing as low epiphytes 


or on rocks in shade or part shade. Abundant on the rocky, scrubby summit of 
Batu Tapah. 


Lindsaya lucida Bl., Enum. Pl. Jav. (1828) 216; Holtt., Gard. Bull. S.S. 9: 131. 
1937, Fl. Mal. 2: 328. 1966. 
L. lobbiana Hk. Sp. Fil. 1: 205. 1848; C. Chr., Gard. Bull. S.S, 4: 396, 
1929; Holtt., Gard. Bull. S.S. 5:61. 1930. 


In Malaya, a streamside plant of low country, widespread. Recorded once 
from the base of a limestone hill growing on limestone boulders by a stream. 


Microlepia speluncae (L.) Moore, var. villossima C. Chr., Gard. Bull. S.S. 4 : 399. 
1929; Holtt., Fl. Mal. 2: 314. 1966. 


Nephrolepis biserrata (Sw.) Schott, Gen. Fil. (1834) t. 3; Holtt., Fl. Mal. 2 : 380. 
1966. 


Aspidium biserratum Sw., Schrad. Jour. 1800/2 : 32. 1801. 


A very common fern of open or partly shaded places. Recorded once from 
limestone (on Gua Musang) as a secondary element about a year and a half after 
fire destroyed the original vegetation. 


Nephrolepis dicksonioides Chr., Verh. Nat. Ges. Basel 11 : 241. 1895; Holtt., Fl. 
Mal. 2 : 376. 1966. 


Distributed in Celebes, Borneo and New Guinea. In Malaya it is restricted to 
limestone, on exposed or partly exposed situations. Common on many hills and 
often forming dense thickets. 


Nephrolepis falcata (Cav.) C.Chr., Dansk. Bot. Ark. 9: 15. 1937; Holtt., Fl. Mal. 
2:381. 1966. 
Tectaria are Cav., Descr. (1802) 250. 
N. barbata Copel., in Holtt., Gard. Bull. S.S. 9 : 132. 1937. 


A common fern on rocky places and as epiphytes. Sometimes on limestone. 
Nephrolepis hirsutula (Forst.) Pr., Tent. Pterid. (1836) 79; Holtt., Fl. Mal. 2 : 382. 
1966. 


Polypodium hirsutulum Forst., Prodr. (1786) 81. 


Common in Malaya in open places. Recorded once from the disturbed base 
of limestone in secondary scrub vegetation. 


Nephrolepis radicans (Burm.) Kuhn, Ann. Lugd. Bat. 4: 285. 1869; Holtt., FI. 
Mal. 2 : 381. 1966. 
Polypodium radicans Burm., FI. Ind. (1768) 233, t. 66. 

Oleandra undulata (Willd.) Ching, Lingnan Sc. Jour. 12: 565. 1933; Holtt., FI. 
Mal. 2 : 384. 1966. 
Polypodium undulatum Willd., Sp. Pl. 5 : 155. 1810. 
A mountain fern, common in Malaya above 1000 m., often on acid peat and 


humus. An unlikely species to be found on limestone. However, one specimen 
(Kadim K491) positively of this, is labelled as from Gunong Tempurong, in forest 


Malayan Limestone Flora 213 


at 250-550 m. The main range of mountains which rises abruptly to over 650 m 
just over a mile to the east of the Gunong Tempurong massive could provide the 
source for the spread of this species. The plant was probably growing on an 
accumulation of humus over the limestone. 


Polystichum lindsaeifolium Ridl., J. Mal. Br. R. As. Soc. 4:61 1926; C. Chr., 


Gard. Bull. S.S. 4 : 393. 1929; Holtt., Fl. Mal. 2: 489. 1966. 
Endemic to limestone in Selangor, Pahang, Kelantan and Perak, uncommon. 


Pteridium aquilinum (L.) Kuhn, var. wightianum (Ag.) Tryon, Rhodora 43 : 1-70. 
1941; Holtt., Fl. Mal. 2 : 634. 1966. 


Pteris aquilina L., Sp. Pl. (1753) 1075. 
Pteridium aquilinum (L.) Kuhn, in Holltt., lc. 389. 
Recorded from limestone as a secondary element. 


Pteridium caudatum (L.) Maxon var. yarrabense Domin, in Tryon, Rhodora 
43 : 63. 1941; Holtt., Fl. Mal. 2: 634. 1966. 
Pteris esculenta Forst., Pl. Escul. (1786) 74. 
Pteridium esculentum (Forst.) Nakai, Bot. Mag. Tokyo 39: 109. 1925; Holtt., 
lc. 390. 


Recorded on limestone as an element of secondary vegetation. 


Pteridrys syrmatica (Willd.) C. Chr. et Ching, Bull. Fan Mem. Inst. Bot. 5: 131. 
1934; Holtt., Fl. Mal. 2 : 530. 1966. 


Aspidium syrmaticum Willd., Sp. Pl. 5 : 277. 1810. 
Distributed in Ceylon, Thailand, Indochina and Malesia to the Philippines. 


In Malaya, this fern is found mainly in the north and nearly always from limestone. 
It is a rock fern of shady places. 


Pteris ensiformis Burm., Fl. Ind. (1768) 230. Holtt., Fl., 2 : 399. 1966. 
Sometimes on limestone. 


Pteris longipinnula Wall., Cat. (1829) 108; Holtt., Fl. Mal. 2: 404. 1966. var. 
b, in Holtt., l.c. 405. 


This variety is endemic to Malaya and chiefly from limestone, though not 
exclusively so. The typical variety is distributed from India to China and south- 
wards throughout Malesia. It differs from the typical variety by the narrower 
pinnae and by the presence of the branched basal pair of pinnae. 


Pteris mertensioides Willd., Sp. Pl. 5: 394. 1810; Holtt., Fl. 2: 404. 1966. 
Pteris scabripes Wall. apud Hook., Sp. Fil. 2: 165. 1858; Holtt., Fl. Mal. 2: 399. 
1966. 


Endemic, usually on rocks by streams, also in lowland forest and rarely on 
limestone, not a common species. 


Pteris tripartita Sw., Sshrad. Jour. 1800/2 : 67. 1801; Holtt., Fl. Mal. 2: 408 1966. 
P. marginata Bory, Voy. 2 : 192. 1804. 
Pteris vittata L., Sp. Pl. (1753) 1074; Hoiltt., Fl. Mal. 2 : 396. 1966. 


Not uncommon on disturbed localities on limestone, by quarry edges and on 
disturbed summits. 


214 Gardens’ Bulletin, Singapore — XXX (1977) 


Tectaria amplifolia (v.A.v.R) C. Chr., Ind. Fil. Suppl. 3: 176. 1934; Holtt., FA. 
Mal. 2: 515. 1966. 


Aspidium amplifolium v.A.v.R., Bull. Jard. Bot. Buit. Ser. II, 11:2. 1913. 


Distributed in Sumatra. Common on and restricted to limestone in Malaya. 
Apparently unknown from the extreme North-west. A plant of shady places. 


Tectaria barberi (Hk.) Copel., Philip. J. Sc. 2c : 414. 1907; Holtt., Fl, Mal. 2 : 508. 
1966. 


Polypodium barberi Hk., Sp. Fil. 5 : 100. 1864. 
Tectaria devexa (Kze) Copel., Philip. J. Sc. 2c: 415. 1907; Holtt., Fl. Mal. 2 : 505. 
1966. 


Aspidium devexum Kze, Bot. Zeit. (1848) 259. 
A. membranaceum Hk., Sp. Fil. 5: 105. 1864. 


Distributed in Ceylon, Thailand and S. China. Restricted to limestone in 
Malaya, widely distributed and fairly common. 


Tectaria griffithii (Bak.) C.Chr., Ind. Fil. (1867) 300; Holtt., Fl. Mal. 2 : 636 1966. 
Nephrodium griffithit Bak., Syn. Fil. (1867) 300. 


Tectaria multicaudata Ching, Sinensia 2: 20. 1931; Holtt., F. Mal. 2: 507. 
1966. 


T. malayense Christ, Philip. J. Sc. 2c : 187. 1907. 


Tectaria macrodonta (Fee) C. Chr., Ind. Fil. Suppl. 3 : 181. 1934; Holtt., Fl. Mal. 
2: 505. 1966. 


Sagenia macrodonta Fée, Gen. Fil. (1852) 313 t. 24. 


A number of specimens collected from north Malaya are doubtfully identified 
as this species. One of these is from limestone. They are smaller than the typical 
form of this species which are represented in India. They resemble T. variolosa 
but the fronds are hairy on both surfaces and hardly dimorphic, and the sori are 
not all on free veins. 


Tectaria variolosa (Wall.) C. Chr., Contr. U.S. Nat. Herb. 26: 289. 1931; Holtt., 
Fl. Mal. 2: 506. 1966. 


Aspidium variolosum Wall., Hook., Spec. Fil. 4: 51. 1862. 


Grammitidaceae 


Ctenopteris alata (Bl.) Holtt., Fl. Mal. 2 : 232. 1966. 
Davallia alata Bl., Enum. Pl. Jav. (1828) 230. 
Prosaptia alata Christ, Ann. Buit. II, 5 : 127. 1905. 


Ctenopteris moultoni (Copel.) C. Chr. et Tard. Not. Syst. 8: 181. 1939; Holtt., Fi. 
Mal. 2 : 229. 1966. 


Polypodium moultoni Copel., Philip. J. Sc. 10c : 149. 1915. 


Hymenophyllaceae 


Trichomanes bipunctatum Poir., in Lamk., Encyl. 8: 69. 1808; Holtt., Fl. Mal. 
Zz: 99. 1966. 


Crepidomanes bipunctatum Copel., Phil. J. Sc. 67: 59. 1938. 


The only common species of this family (except in the very north) on limestone. 


Malayan Limestone Flora 215 


Trichomanes christii Copel., Phil. J. Sc. 1. Suppl. 251. 1906, ibid., 51 : 185. 1933; 
Holtt., Fl. Mal. 2: 100. 1966. 


Crepidomanes christii Copel., Phil. J. Sc. 67 : 60. 1938. 

Trichomanes humile Forst., Prodr., (1786) 84. Copel., Phil. J. Sc. 51: 164. 1933; 
Holtt., Fl. Mal. 2: 98. 1966. 
Crepidopteris humilis Copel., Phil. J. Sc. 67 : 58. 1938. 


Trichomanes motleyi Bosch, Ned. Kruidk. Arch. 5: 145. 1861; Copel., Phil. J. Sc. 
51:201. 1933; Holtt., Fl. Mal. 2:92. 1966. 


Polypodiaceae 
Crypsinus enervis (Cav.) Copel., Gen. Fil. (1947) 207; Holtt., Fl. Mal. 2: 199. 
1966. 


Polypodium enervis Cav., Descr. (1802) 245. 


Drynaria bonii Christ., Not. Syst. 1 : 186. 1910; Tardieu-Blot et Christ., Fl. Gen. 
Indochine 7 : 515. 1941. 


Rhizome creeping. Nest leaves small, ovate, base cordate-sagittate, margin 
entire scalloped or shallowly lobed, 5-10 by 3.5—6.5 cm. Foliage leaves with stipes 
8-20 cm long, narrowly winged; lamina deeply lobed, 20-45 by 12-20 cm. Lobes 
3-7 pairs, 8-20 by 2-4 cm, base decurrent along the rachis. Sori non-indusiate and 
small, scattered on the lower surface of frond. 


Distributed in S. China, Indochina and Thailand. A new record (Chin 1764) 
for Malaya. Common as low epiphytes and on rocks in Pulau Langgun, Langkawi 
on the northern side of the island. 


Characterised by the small unlobed nest leaves and the dissection of the 
foliage leaves. The old rhizome which is fleshy and flattened grows up to 3 by 
1.5 cm. 


Drynaria quercifolia (L.) J. Sm., J. Bot. 3: 398. 1841; Holtt., Fl. Mal. 2: 182. 
1966. 


Polypodium quercifolium Linn., Sp. Pl. (1753) 1087. 


Drynaria rigidula (Sw.) Bedd., Ferns Brit. Ind. (1869) t. 314; Holtt., Fl. Mal. 
2: 183. 1966. : 


Polypodium rigidulum Sw., Schrad. J. 1800/2 : 26. 1801. 


A common epiphyte in the northern half of Malaya; it is not found in the 
south. It is not uncommon on limestone in Langkawi and has been recorded as 
far south as Bukit Takun in Selangor. This forms the southernmost record for this 


Poe in Malaya. On limestone it has been recorded as epiphytes as well as on 
rocks. 


eta sparsisora (Desv.) Moore, Ind. Fil. (1862) 348; Holtt., Fl. Mal. 2: 183. 


Polypodium sparsisorum Desv., Berl. Mag. 5 : 315. 1811. 
Lemmaphyllum accedens (Bl.) Donk. Reinw. 2 : 409. 1954; Holtt., Fl. Mal. 2: 152. 
1966. 


Polypodium accedens Bl., Enum. Pl. Jav. (1828) 121. 
Weatherbya accedens Copel., Gen. Fil. (1947) 191. 
Lepisorus longifolius (Bl.) Holtt., Fl. Mal. 2: 151. 1966. 
Grammitis longifolia Bl., Enum. Pl. Jav. (1828) 119. 
Paragramma longifolia Moore, Copel., Gen. Fil. (1947) 190. 


216 Gardens’ Bulletin, Singapore — XXX (1977) 


Leptochilus decurrens Bl., Enum. Pl. Jav. (1828) 206; Holtt., FI. va 2: 164 1966. 
Acrostichum variabile Hook, Sp. Fil. 5 : 277. 1864. 

Perper avenia (BIl.) Presl., Tent. Pterid. (1836) 215; Holtt., Fl. Mal. 2 : 167. 
Grammitis avenia Bl., Enum. Pl. Jav. (1828) 117. 
Loxogramme blumeana Presl., Tent. Pterid. (1836) 215. 
Not common on limestone, but recently found to be abundant on the rocky, 


one-layered scrub forest on the summit of Gua Batu Boh, near Gua Musang, 
Kelantan; on rocks and as low epiphytes. 


Loxogramme scolopendrina (Bory) Presl., Tent. Pterid. (1836) 215; Holtt., Fl. Mal. 
2: 168. 1966. 
Grammitis scolopendrina Bory, Dup. Voy. (1829) 257. 
Fairly common on limestone and widely distributed. 

Microsorium musifolium (Bl.) Ching, Bull. Fan Mem. Inst. 4: 295. 1933; Holtt., 
Fl. Mal. 2: 176. 1966. 
Polypodium musifolium Bl., Enum. Pl. Jav. (1828) 134. 

Microsorium punctatum (L.) Copel., Univ. Cal. Publ. Bot. 16: 111. 1929; Holtt., 
Fl. Mal. 2: 179. 1966. 
Acrostichum punctatum Linn., Sp. Pl. Ed. 2. (1763) 1524. 


Photinopteris speciosa (BL.) Pres]., Epim. Bot. (1849) 264; Holtt., Fl: Mal. 2 : 187. 
1966. | 


Lomaria speciosa Bl\., Enum. Pl. Jav. (1828) 202. 
Photinopteris rigida Bedd., Fl. Brit. Ind. (1867) t. 211. 
Phymatodes nigrescens (Bl.) J. Sm., Ferns Br. & For. (1866) 94; Holtt., Fl. Mal. 
2: 193. 1966. 
Polypodium nigrescens P\., Enum. Pl. Jav. (1828) 126. 
Phymatodes scolopendria (Burm.) Ching, Contr. Inst. Bot. Nat. Acad. Peip. 2 : 63. 
1933; Holtt., Fl. Mal. 2: 191. 1966. 
Polypodium scolopendria Burm., Fl. Ind. (1769) 232. 
P. phymatodes Linn., Mant. (1771) 306. 


Recorded a number of times from limestone, ahatiddte on Batu Tapah. 
Kelantan, on the scrubby summit growing over boulders and as low epiphytes. 


Polypodium papillosum Bl., Enum. Pl. Jav. (1828) 131; Holtt., Fl. Mal. 2: 203. 
1966. 


Pyrrosia adnascens (Sw.) Ching, Bull. Chin. Bot. Soc. 1 : 45. 1935; Holtt., Fl. Mal. 
2: 144. 1966. 7 


Polypodium adnascens Sw., Syn. Fil. (1806) 25. 


Common in the lowlands of Malaya, epiphytic pres on Tocks; also common on 
limestone and widely collected. 


Pyrrosia floccigera (Bl.) Ciuag, Bull. Chin. Bot. Soc. 1: 71. 1935; Holtt., Fl. Mal. 
2: 147. 1966. 


Niphobolus flocciger Bl., Enum. Pl. Jav. (1828) 107. 


Malayan Limestone Flora 217 


Fairly common in Malaya, usually from 300-1300 m, in partly shaded areas, 
epiphytic. Recorded once from limestone at less than 100 m elevation (UNESCO 
1962, 216, from Gua Batu Boh, Kelantan). 


Pyrrosia penangiana (Hook.) Holtt., Fl. Mal. 2: 146. 1966. 
Niphobolus penangianus Hook., Ic. Pl. (1840) t. 203. 


Distributed in Sumatra. Recorded from the northern half of Malaya, usually 
on limestone; on rocks or as epiphytes. 


Pyrrosia stigmosa (Sw.) Ching, Bull. Chin. Bot. Soc. 1:67. 1935; Holtt., Fl. Mal. 
2: 148. 1966. 
Polypodium stigmosum Sw., Schrad. Jour. 1800/2: 21. 1801. 
Distributed in North India and Indochina and from Sumatra to New Guinea. 


Restricted to limestone in Malaya, on rocks and epiphytic; it has been reported 
as an epiphyte on Rain trees near Gua Batu, Selangor. 


Pyrrosia varia (Kaulf.) Farwell, Am. Midl. Nat. 12 : 302. 1931; Holtt., Fl. Mal. 
2: 146. 1966. 
Niphobolus varius Kaulf., Enum. Fil. (1824) 125. 


A widely distributed species, also common on limestone. 


Schizaeaceae 


Lygodium flexuosum (L.) Sw., Schrad. Jour. 1800/2: 106. 1801; Holtt.. Fl. Mal. 
Zt. 1966. 


Ophioglossum flexuosum Linn., Sp. Pl. (1753) 1063. 


Lygodium polystachyum Wall. ex Moore, Gard. Chron. (1859) 671; Holtt., Fl. Mal. 
2:56. 1966. 


Distributed in Burma and Thailand. Uncommon and apart from specimens 
from Penang Hill (granite), this species is restricted to limestone in Malaya. 
Recorded from Perak, Kelantan, Langkawi and Pahang, usually in shady forest. 


Schizaea inopinata Selling, Sevensk Bot. Tidskr. 40 : 274. 1946; Holtt., Fl. Mal. 
2252. 1966. 


Distributed in Sumatra and Philippines. In Malaya, this species is restricted 
to limestone in Kelantan, Pahang and Selangor, not uncommon. It resembles the 
common S. digitata (L.) Sw. and is frequently mistaken for it. S. inopinata is how- 
ever characterized by having the sporangia in two instead of four rows. 


Thelypteridaceae 


Abacopteris urophylla (Wall.) Ching, Bull. Fan Mem. Inst. Bot. 8: 251. 1938; 
Holtt., Fl. Mal. 2: 296 1966. 


Polypodium urophyllum Wall., in Hook., Sp. Fil. 5: 9. 1863. 

Cyclosorus extensus (Bl.) Ching, Bull. Fan Mem. Inst. Bot. 8 : 182. 7938: Holtt., 
Fl. Mal. 2: 264. 1966. 
Aspidium extensum Bl., Enum. Pl. Jav. (1828) 156. 

Cyclosorus interruptus (Willd.) Ching, Bull. Fan Mem. Inst. Bot. 8 : 184. 1938; 
Holtt., Fl. Mal. 2 : 262. 1966. 
Pteris interrupta Willd., Phytogr. 1 : 13. 1794. 


218 Gardens’ Bulletin, Singapore — XXX (1977) 


Cyclosorus megaphyllus (Mett.) Ching, Bull. Fan Mem. Inst. Bot. 8: 225. 1938; 
Holtt., Fl. Mal. 2: 268. 1966. 


Aspidium megaphyllum Mett., Ann. Mus. Lugd. Bat. 1 : 233. 1864. 
Cyclosorus unitus (L.) Ching, Bull. Fan Mem. Inst. Bot. 8: 192. 1938; Holtt., FI. 
Mal. 2: 260. 1966. 
Polypodium unitum L., Syst. Nat. ed. 10, 2 : 1326. 1759. 
Thelypteris immersa (Bl.) Ching, Bull. Fan Mem. Inst. Bot. 6 : 306. 1936; Holtt., 
FL 2; 245.. 1906. 
Aspidium immersum Bl., Enum. (1828) 156. 


GYMNOSPERMS 
Araucariaceae 


Agathis dammara (Lambert) L. G. Rich., Comm. Bot. Conif. Cycad. (1826) 83: 
Keng, Tree Fl. Mal. 1:41. 1972. 


A. loranthifolia Salisb. in Ridl., Flora 5 : 278. 1925. 
A. alba (Lam.) Jeff. in Burk., Dict. I: 62. 1935. Corner, Ways. Trees I: 715. 
1940. 


Usually found on the hills; once recorded (Loh FRI 17201) for limestone 
at 600-700 m at Gua Peringat, Pahang. This was growing in deep soil in a gully. 


Cycadaceae 


Leaves with petioles about 100 cm long, leaflets 0.7-0.9 cm wide. Male cone 
about 20 cm) long >) 407-24.:.c0d dodge aes lal aren Cycas siamensis 
Leaves with petioles about 200 cm long, leaflets 1.3-2.0 cm wide. Male cone 
about 35 cm lome yi peccdskpcectabn anee ee tk eee aaa on Saeed Cycas rumphii 


Cycas rumphii Mig., Bull. Sc. Phys. Neerl. 839, 45; Hk. f., F.B.I. 5: 657; Ridl.. 
Fl. 5 : 284. 1925. 


Cycas siamensis Mig., Bot Zeit. (1863) 334; Hk. f., F.B.I. 5 : 657; Ridl. Fl. 5 : 285. 
1925. 


Distributed in Burma and Thailand; in Malaya, restricted to limestone, and 
common on Langkawi islands. 


Gnetaceae 


A shrub or small tree, to 3 m tall; flowers not embedded in thick hair masses. 
ee et ee eee Pe oe ee Gnetum gnemon var. tenerum 
A liana, more than 10 m long; flowers embedded in thick hair masses. ............... 
a vos Gnd cs'ovwavan dulro0 0 bis apmpondin anc pl a ad i eepelccadio i a ne Gnetum cuspidatum 


Gnetum cuspidatum Bl., Rumphia 4:5. 1848; Henders., J. Mal. Br. As. Soc. — 
17 : 87. 1939; Markgraf, Fl. Mal. I, 4: 343. 1951. 
G. penangense Ridl., Fl. 5 : 276. 1925. 

Gnetum gnemon Linn., Mant. 1: 125. 1767; Markgraf, Bull. Jard. Bot. Btzg II, 
(10) : 436. 1930; Rid]. Fl. 5:273. 1925; Markgraf, Fl. Mal. I, 4: 340. 1951. 
var. tenerum Markgraf, Fl. Mal. I, 4: 341. 1951. 


Malayan Limestone Flora 219 


Podocarpaceae 


i A en Se Se ee ee ee Podocar pus polystachyus 
Male cones solitary or sometimes 2-3 together. Leaves usually larger, 5.5—20 
i I Ee on eee A ae OA Ae Podocar pus _neriifolius 


Podocar pus polystachyus R. Br. [in Mirb., Mem. Mus. Hist. nat. Paris 13:47. 1825 
nom. nud.] ex Endl., syn. Conif. (1847) 215. Ridl., Fl. 5: 282. 1925; Keng, 
Ttee Fl, Mal, 1: 53: 1972. 


Usually coastal except on limestone hills where it is very widely distributed 
and common. 
Podocar pus neritifolius D. Don, in Lambert, Desc. Gen. Pinus 1 : 21. 1824; Ridl., FI. 
5 : 281. 1825; Keng, Tree Fl. Mal. 1 : 51. 1972. 


Usually on hill and mountain forest, rarely below 300 m. Recorded on lime- 
stone from Gua Batu Selangor. (Ng FRI. 1634). 


oe 


. ‘ } ; ¢ ba | ¥ 
Vi Pre Le . etal 
om Yarend pitsp ener seek e a 
: & 4 A ‘ 
‘ j HI 6% 


aS eats “4 saa * . 
sr a ay - 7: coms tent Natt oe 


CBIR et haben saa 


ano ;eSel Se: 2 nig, Pig, Ea cin ag 


betudrnietb ¥ lobivw Woy, aoe HAY, aig. arene nO br i 


MM bit C8! 10: 1 eunit.0sd_o7e) unedited) Al nom ax a “at 

crop fe ot Vee 4 ome a 

nl of bsbhio: SE ct (KH solse nA Le mrss fa): rrisSrtapcigay bne Hid 
. {BEDE (34 3) sognalee viel 


+ ee 
"ik 1 Pais & | 
“ ‘Sa 
~— 
: ts 
* 
c 5 yr; -% 
4 
Litas. OF 


aT menace ree 


pity 


A List of Holttum’s New Taxa and Name-Changes in Ferns to 
July 1975 


by 
J. A. CRABBE 


British Museum (Natural History) 


London 


Little did I think, when I embarked on this abstracting, that I would be 
delving among some 420 titles by R. E. Holttum which range over botany, 
horticulture, philosophy, etc. (see Flora Malesiana Bulletin 28: 1975), and that 
IT would find around 700 items for this present list. 


I have not included anything published after his 80th birthday on 20 July 
1975, nor some few names superfluously replicated, nor some invalidly published 
(usually without latin diagnoses and usually rectified later). The genus Polystic- 
hopsis (17: 149, 152) is, to me, obscure and I suspect there may be other items in 
this work and also in a similar treatment (The classification of ferns, in Biol. 
Rev. 24: 267-296. 1949). 


It is with the greatest pleasure that I present this list, acknowledging with 
gratitude much helpful discussion with colleagues, and trusting it will serve a useful 
purpose. 

For brevity, I have numbered the bibliography, then quoted those numbers 
followed by page numbers, New species give the type and its location, a semicolon 
separating holotype from isotypes. Where Holttum designates no type but lists 
specimens he has seen, I have quoted the first in that list, not thereby implying 
nid other status. Basionym authors followed by their genus are enclosed in round 

rackets. 


Abacopteris menisciicarpa (Bl. — Aspidium) 24:290 
A, peltochlamys (C. Chr. — Dryopteris) 24:295 
A, rubicunda (v.A.v.R. — Phegopteris) 24:292 
A. salicifolia (Wall. ex Hook. — Meniscium) 24:288 


Alsophila parvifolia 2:19. — Mentawi Is., Kloss 14579 (SING) 


Amauropelta bergiana (Schlechtend. — Polypodium) 57:133 

. bergiana var, calva 57:133.— Cameroon Mt, Richards 4308 (K) 

. bergiana var. tristanensis 57:134. — Tristan da Cunha, Carmichael (K) 
. heteroptera (Desv. — Nephrodium) 57:132 

. membranifera (C, Chr. — Dryopteris) 57:132 

. oppositiformis (C. Chr. — Dryopteris) 57:135 

. Strigosa (Willd. — Aspidium) 57:134 

. tomentosa (Thouars — Polypodium) 57: 134 


AMPHINEURON gen. nov. 50:45. figs 19, 19a. — type species: A. opulentum 
A. immersum (Blume — Aspidium) 62:203 

A. opulentum (Kaulf.— Aspidium) 50:45 

A. terminans (Hook: — Nephrodium) 54:82 


>Prr>rrprpyp 


221 


222 Gardens’ Bulletin, Singapore — XXX (1977) 


Araiostegia membranulosa (Wall. ex Hook. — Davallia) 52:230 


Arcypteris brongniartii (Bory — Polypodium) 21:195 
A. gigantea (Cesati— Nephrodium) 21:195 

A. irregularis (Presl — Polypodium) 21:193. ff. 1-3 
A. macrodonta (Fée — Dictyopteris) 21:194 


Arthropteris wollastonii (Ridl.— Polypodium) 39:229 
Aspidium maingayi (Bak.— Gymnogramine) 5: 207. ff. 1-9 


Asplenium kinabaluense 7:281, t. 62 — Borneo, Clemens 29830 (SING) 
A. malayo-alpinum 7:280. t. 61 — Borneo, Holttum 25485 (SING) 
A. nidus var. phyllitidis (Don pro spec.) — 2:20 


Athyrium amplissimum (Bak. — Asplenium) 24:569 

A. angustipinna 23:8 — Malaya, Holttum SFN 39461 (SING) 
. angustisquamatum 16:273.— Malaya, Holttum SFN 37359 (SING) 
. curtisii 22:545. — Malaya, Molesworth-Allen 2338 (SING) 
. insigne (Holtt.— Diplazium) 24:562 

. kunstleri (Holtt. — Diplazium) 24:564 

latisquamatum (Holtt.— Diplazium) 24:563 

malaccense (Presl.— Diplazium) 24:552 

montanum (v.A.v.R. — Diplazium) 24:555 

. prescottianum (Wall. ex Hook. — Asplenium) 24:557 

. procumbens (Holtt. — Diplazium) 24:572 

. Tiparium (Holtt.— Diplazium) 24:554 

. Simplicivenium (Holtt.— Diplazium) 24:573 

. subintegrum (Holtt.— Diplazium) 24:557 

. velutinum (Holtt. — Diplazium 24:567 


Belvisia longissima 41:328.— New Guinea, Pullen 5356 (K) 


Bolbitis nitens 27:453. — Malay Peninsula, Molesworth-Allen 3384 (K) 
B. singaporensis 16:271.— Malaya, Hullett s.n., March 1882, 2 sheets (SING) 


Cerosora sumatrana 27:450.— Sumatra, Meijer 3959 (L) 


CHINGIA gen. nov. 50:31. — type species: C. ferox 

. acutidens 60:17. — Celebes, Alston 15708 (BM) 

. atrospinosa (C. Chr. — Dryopteris) 60:19 

. Clavipilosa 60:23. N Borneo, Mt Kinabalu, Holttum 44 (SING; K) 

christii (Copel. — Cyclosorus) 60:25 

ferox (Bl. — Aspidium) 50:31 

horridipes (v.A.v.R. — Dryopteris) 60:26 

imponens (Cesati— Polypodium) 60:21 

malodora (Copel. — Dryopteris) 60:19 

. longissima (Brack. — Goniopteris) 60:17 

. mMuricata (Brause — Dryopteris) 60:20 

. paucipaleata 60:26. — Philippine Is., Luzon, Price 781 (PNH; K) 

. perrigida (v.A.v.R. — Phegopteris) 60:27 

. Supraspinigera (Rosenst. — Dryopteris) 60:27 

. pricei 60:22. — Philippine Is., Luzon, Price 512 (PNH; K) 3 
. pseudoferox 60:24. — Malay Peninsula, Perak, Matthew s.n., 6 Feb 1908 (K) 
. Sambasensis 60:24. W Borneo, Brooks s.n, Sept 1908 (BM; K, L) 

. urens 60:28. — Philippine Is., Luzon, Mt Makiling, Price 2367 (PNH; K) 


Christella sect. PELAZONEURON 57:144 — type species: Polypodium patens 
C. afzelii (C. Chr. — Dryopteris) 57:146 
C. altissima 57:141.— Natal, Buchanan 103b (K) 


> >>> > DP DL DD SD 


anaaAANAAARAAAAAANANA 


Fern names of R. E. Holttum 223 


(Christella cont.) 


anaaaaaaaaaaanaanaan ananaanann 


. arida (D,. Don — Aspidium) 62:206 

. callensii (Alston — Cyclosorus) 57:146 

. assamica (Bedd.-— Nephrodium multilineatum var.) 62:210 
. chaseana (Schelpe — Thelypteris) 57:148 

. Clarkei (Bedd. — Pleocnemia) 62:203 


crinipes (Hook. — Nephrodium) 62:208 

cylindrothrix (Rosenst.— Dryopteris) 62:208 

dentata (Forsk. — Polypodium) 55:306, [transfer previously made by Brownsey 
& Jermy in Brit. Fern Gaz. 10:338. 1973] 

distans (Hook. — Nephrodium) 57:140 


. evoluta (Clarke & Baker — Nephrodium amboinense var.) 62:208 


friesii (Brause — Dryopteris) 57:145 


. gueinziana (Mett.— Aspidium) 57:147 
. guineensis (Christ — Dryopteris) 57:145 


hilsenbergii (Pres! — Nephrodium) 57:142 

jerdonii (Ching — Cyclosorus) 62:206 

lebeufii (“‘leboefii’?) (Baker — Polypodium) 62:206 
meeboldii (Rosenst.— Dryopteris) 62:208 
microbasis (Baker — Nephrodium) 57:146 
modesta 57:144. — Madagascar, Decary 16,485 (P) 


. molliuscula (Kuhn — Aspidium) 62:205 
. multifrons (C. Chr. — Dryopteris) 57:145 


namburensis (Bedd. — Polypodium) 62:206 


. papilio (Hope — Nephrodium) 62:208 

. papyracea (Bedd.— Nephrodium) 62:206 
. prolixa (Willd. — Aspidium) 54:83 

. zeylanica (Fée — Nephrodium) 62:208 


Cibotium arachnoideum (C. Chr. — Cibotium cumingii var.) 32:166 


Colysis acuminata (Bak. — Gymnogramme) 24:162 
CORYPHOPTERIS gen. nov. 50:33. — type species C. viscosa 


4 
c 


didymochlaenoides (Clarke — Nephrodium gracilescens var.) 62:203 
hirsutipes (Clarke — Nephrodium gracilescens var.) 62:203 
viscosa (Bak. — Nephrodium) 50:33 


Crypsinus laciniatus (Presl — Phymatodes) 24:198 


& 
3d 


stenophyllus (Bl. — Polypodium) 24:199 
subfasciatus (Holtt.— Phymatodes) 24:202 


Ctenitis manilensis (Presl. — Lastrea) 47:36 


ad 


setosa (Presl — Lastrea) 47:20 


Ctenopteris alata (Bl. — Davallia) 24:232 


ANAAANANAAN 


. brevivenosa (v.A.v.R. — Polypodium) 24:228 
. contigua (Forst. — Trichomanes) 24:230 


khasyana (Hook. — Polypodium) 24:233 


. leysii (Bak. — Polypodium) 24:232 

. minuta (Bl. — Polypodium) 24:228 

. moultonii (Copel. — Polypodium) 24:229 

. obliquata (Bl. — Polypodium) 24:229 

. Sparsipilosa 16:268.— Malaya, Holttum SFN 23522 (SING) 
. Subminuta (v.A.v.R. — Polypodium) 24:228 


Cyathea subgen. Cyathea sect. GYMNOSPHAERA (BI. — pro gen.) 32:115 
C. subgen. SPHAEROPTERIS (Bernh. — pro gen.) 32:124 


C. 


subgen. Sphaeropteris sect. SCHIZOCAENA (J. Sm. — pro gen.) 32:141 


224 Gardens’ Bulletin, Singapore — XXX (1977) 


(Cyathea cont.) 


fepeteerne 2 MM NANNNAAANANAANAANAANAAN ANNHAAAN AANAARA AN, Gee 


subgen. Sphaeropteris sect, Schizocaena subsect. SARCOPHOLIS 32: 153. fig. 30. 
Type species: Cyathea rosenstockii — 

subgen, Sphaeropteris sect. Sphaeropteris subsect. FOURNIERA (Bommer — 
pro gen.) 32:138. fig. 27 


. acanthophora 30:51.—N. Borneo, Clemens 34012 (BO; US) 


aeneifolia var. macrophylla 32:134.—-NE New Guinea, Hoogland & Schodde 
7209 (L) 


. aeneifolia var, melanacantha (Copel. — Cyathea) 32:134 
. agathetii 30:51. — Indonesian Borneo, Kostermans 12870 (BO; A, K, L) 
. alleniae 30:52. — Pahang, Molesworth Allen 4127 (K; US) 


angustipinna 30:52. — Sarawak, Richards 1675 (K; SING, US) 
archboldii var, horrida 32:99. NE New Guinea, Hoogland & Pullen 5506 (K) 


. atrospinosa 30:53. —- NE New Guinea, Hoogland & Pullen 6090 (K) 


atrox var. inermis 32: 130. figs. 24, 25. — NE New Guinea, Hoogland & Schodde 
7457 (L) 


. australis subsp. norfolkensis 33:244. — Norfolk Id., Metcalf s.n. (NSW, P5817) 
. carrii 30:53. — Papua, Carr 13526 (K; BM, L) 

. catillifera 30:53. Papua, Brass 4549 (BRI) 

. cicatricosa 33:274. — New Caledonia, Green 1778 (K) 

. coactilis 31:533.— Papua, Schodde 1887 (CANB) 


cucullifera 30:54. —- NE New Guinea, Hoogland & Pullen 5497 (K; BO, BRI, 
L, SY, Us) 

decurrens subsp. epaleata 33:249. — Tahiti, Grant 4233 (K) 
deminuens nom, nov. 32:145.— Alsophila parvifolia 

dicksonioides 31:529.—- New Guinea, Hoogland & Schodde 7171 (L) 

discophora 30:54. N. Borneo, Clemens 31698 (B; BO, US) 

eriophora 30:55. — Papua, Carr 14439 (K; BM, L) 

excavata 8:306, t.31. — Malaya, Holttum SFN 23538 (SING) 

gigantea (Wall. — Alsophila) 8:318 

glaberrima 30:55. — Papua, Brass 27092 (L; US) 

hooglandii 30:56. -NE New Guinea, Hoogland & Schodde 7203 (L) 

inaequalis 30:56. — Papua, Brass 23547 (L; A) 

insulana 30:56. — Papua, Brass 24725 (A; L) 

insularum 30:57. — Papua, Brass 27419 (L; K, US) 

jacobsii 57:499.— S. Sumatra, Jacobs 8113 (L; BO, K) 

kanehirae nom. nov. 32:113.— Alsophila arfakensis 

latebrosa var, indusiata 8:305.— Malaya, Holttum SFN 23539 (SING) 

loerzingii 30:58. — Sumatra, Lérzing 14904 (L; BO) 

lunulata subsp. vitiensis (Carr. — Alsophila vitiensis) 33:264 


. macrophylla var. quadripinnata 32:141.— NE New Guinea, Womersley NGF 


13959 (K) 


. mesosora 30:57. —-_NE New Guinea, Robbins 1032 (L) 
. microchlamys 30:58. — Philippine Is., Ramos 30475 (US; P) 
. nigrolineata 30:58. NE New Guinea, Hoogland & Pullen 5495 (K; BO, BRI, 


L, SYD, US) 


. higropaleata 30:59. NE New Guinea, Pullen 666 (L) 
. nilgirensis 35:468. — India, Gamble 17799 (K) 


nothofagorum 41:327.— NE New Guinea, Pullen 5358 (K) 


. Odsora 30:59.—N. Borneo, Clemens 51188 (K; A, L, MICH, UC) > 

. pallidipaleata 30:60.— Celebes, Eyma 776 (BO) 

. parvipinna 30:60.— Papua, Brass 25837 (L; K, US) 

. pseudomuelleri 30:61. — W. New Guinea, Brass 9430 (A; BO, L, MICH, UC) 
- pycnoneura 31:533.— New Guinea, Pullen 562 (CANB: BM, L) 

. robusta (C. Moore — Alsophila) 33:265 


Fern names of R. E. Holttum 225 


(Cyathea cont.) 


C. rubella 30:61. — Papua, Hoogland 4487 (K; A, BM, BRI, L) 

. Sarasinorum nom, nov. 30:61.— Alsophila contaminans var. longepaleata 

. semiamplectens 30:62.—- NE New Guinea, Womersley 11500 (BRI; L) 

. setifera 30:62. N. Moluccas, Main & Aden 1306 (BO; K, L) 

solomonensis 33:252.— Solomon Is., Whitmore & Womersley BSIP 1035 
(LAE) 

subtripinnata 31:534. — Papua, Schodde 1763 (CANB) 

. stelligera 33:250.— New Caledonia, McKee 9863 (NSW) 

. stipitipinnula 30:62.— N. Borneo, Clemens 33156 (K; A, BO, L, UC) 

. womersleyi 30:63. -NE New Guinea, Womersley & Millar 8470 (K; A, BO. 
L, SYD) 

vandeusenii 31:529.—- New Guinea, Brass 29989 (L; K) 

. verrucosa 30:63. — Sumatra, Matthew s.n. (K) 


AA AA|ANM AAAN 


QO 


yclosorus deminuens 34:135.— New Guinea, Brass 32106 (K) 
dicranogramma (v.A.v.R.— Dryopteris) 24:267 

ecallosus 16:269.— Malaya, Holttum SFN 31294 (SING) 

heterocarpus var, glaucostipes (Bedd.— Nephrodium glaucostipes 24:27] 
. jacobsii 31:530. — Sarawak, Jacobs 5086 (L: K, SAR, G, US, B, S-PA) 
. Menisciicarpus (Blume — Aspidium) 44:39 

. oligodictyus (Bak. — Acrostichum) 31:530 

. polycarpus (Bl.— Aspidium) 24:283 


Davallia dimorpha 9:122.— Malaya, Holttum SFN 31298 (SING) 
D. trichomanoides var. lorrainii (Hance — Davallia lorrainii) 24:361 


Dicksonia lanigera 30:64. W. New Guinea, Pulle 1010 (L; BM) 
D. mollis 30:64. — Philippine Is., Elmer 9874 (K; L, MICH, US) 


Dicranopteris subgen. ACROPTERYGIUM (Diels, pro sect. sub Gleichenia 
subgen. Mertensia 26:261 

. Clemensiae 26:275 —N. Borneo, Clemens 28745 (BM; K, L, US) 

. linearis var. alternans (Mett. — Gleichenia dichotoma var, 26:278 

. linearis var, altissima 26:276.— Malay Peninsula, Corner SF 31447 (SING: 
KL) 

linearis var. demota 26:275 — N. Borneo, Clemens 29535 (BM; K, L, US) 

linearis var. dichotoma (Thunb. — Polypodium dichotomum) 26:277 

linearis var. ferruginea (Blume — Gleichenia ferruginea) 26:278 

linearis var. glabra 44:40— Bismarck Archipelago, Noona Dan Exped. 
2608 (C) 

linearis var. inaequalis (Rosenst. — Gleichenia) 26:278 

linearis var. latiloba 26:277. — Philippine Is., Merril] 975 (US) 

linearis var. montana 26:276.— Malay Peninsula, Molesworth Allen 2720 
(SING) 

linearis var. rigida (Blume — Gleichenia hermannii var.) 26:277 

linearis var. subpectinata (Christ — Gleichenia subpectinata 26:277 

linearis var. subspeciosa 26:278.— N. Borneo, Topping 1516 (SING; US) 

speciosa (Pres]l — Hicriopteris) 26:273 


aAAAANNA 


O 9959 9S9 9995 995 


iplazium angustipinna (Holtt.— Athyrium) 45:637 

armatum (Copel. — Lastrea) 34:135 

curtisii (Holtt.— Athyrium) 45:637 

dielsii (Brause — Alsophila) 34: 136 

. insigne 9:123.— Malaya, Holttum SFN 21635 (SING) 

. kunstleri 15:88.f.2.— Malaya; Holttum SFN 31194 (SING) 

. latilobum 43:308. — Thailand, Hansen, Seidenfaden & Smitinand 11155 (C; K) 
. latisquamatum 9:124.— Malaya, Holttum SFN 31311 (SING) 


whelvieivleic) 


226 Gardens’ Bulletin, Singapore — XXX (1977) 


(Diplazium cont.) 


. procumbens 15:95.f.4. — Malaya, Holttum SFN 36503 (SING) 

. Tiparium 15:97.f.5.— Malaya, Hume 8186 

. simplicivenium 15:100.f.6.— Malaya, Holttum SFN 36507 (SING) 
. Speciosum var. major (Bedd. — D, sorzogonense var.) 15:103 

. stellatopilosum (Brause — Dryopteris) 34: 136 

. Stipitipinnula 41:328 New Guinea, Hoogland 9090 (K) 
subintegrum 9:125.— Malaya, Holttum 31350 (SING) 

. Velutinum 9:126.— Malaya, Holttum SFN 31221 (SING) 

. woodwardioides (Presl — Brachysorus) 47:30 


wluluieieieioioie) 


Diplora translucens 44:41. — Bismarck Archipelago, Noona Dan Exped. 1824 (C) 


Dryopteris brunnea var. glabrata (Clarke — Polypodium distans var.) 9:127 
D. haniffii 16: 272. f. 1.— Malaya, Holttum SFN 31286 (SING) 
D. urophylla var. nitida 7:249, 251 (text-f.) — Borneo, Holttum 25592 (SING) 


Elaphoglossum angustifrons 40:319. NE New Guinea, Womersley 11272 (LAE; 
. apiculatum 40:320.— W. New Guinea, Brass 13624 (G) 

. apoense 40:320. — Philippine Is., Edafio PNH 710 (MICH) 

. arachnoideum 40:320.— Papua, Brass 24541 (A) 

. brevifolium 16:270.— Malaya, Holttum SFN 20750 (SING) 
calanasanicum 64:196. — Philippine Is., Luzon, Price 2933 (K; PNH) 
favigerum 40:321.— W. New Guinea, Brass 13440 (G; MICH) 
heterostipes 40:321.—C. Indonesian Borneo, Endert 4424 (BO) 
idenburgensis 40:321. —- W. New Guinea, Brass 12283 (G; MICH) 
indrapurae 40:321.— Sumatra, Alston 14275 (BM) 

latemarginatum 40:322.— Papua, Brass 24896 (A) 

malayense 40:322.— Malay Peninsula, Henderson 17765 (K; SING) 
melanochlamys 40:322.—-W. New Guinea, Eyma 5408 (BO) 
negrosensis 40:323.— Philippine Is., Elmer 9885 (MICH; BO) 
nesioticum 40:323.— Papua, Brass 24882 (A) 

. nigripes 40:323. — W. New Guinea, Pulle 493 (BM; BO, L, U) 
ophioglossoides (Goldm. — Acrostichum) 64:197 

. pallescens 40:323.— Papua, Brass 22879 (A) 

peninsulare 16:270. — Siam, Eryl Smith 2021 (SING) 

planicosta 40:324.—- NE New Guinea, Jermy 4216 (BM) 

. pullenii 40:324.-_NE New Guinea, Pullen 5034 partim (L) 

. resiniferum 40:324.— Papua, Wakefield 1466 (BM) 

. spongophyllum Bell ex. 40:325.— N. Borneo, Clemens 31869 (BO) 
. Stenolepis Bell ex. 40:325.— N. Borneo, Clemens 28019 (US) 

. Ssumatranum 40:325.— Sumatra, Matthew s.n. (K) 

. thamnopteris 40:326.— Papua, Brass 22990 (A) 

. vepriferum 40:326.—-NE New Guinea, Clemens 7417 (B; G) 


ot oe subgen, DIPLOPTERYGIUM — (Diels, pro sect. sub subgen. Merten- 
sia :261 

. alstonii 26:267.— Sumatra, Alston 14981 (BM) 

. angustiloba 26:263.—N. Guinea, Brass 4960 (BM; K) 

. brevipinnula 26:264.— Sarawak, Bell 2042 (BM) 

clemensiae (Copel. — Hicriopteris) 28:14 

clemensiae var, membranacea (Holtt.— Gleichenia papuana var.) 28:14 
. deflexa 26:264.— Addendum, New Guinea, Papua, Brass 27171 (L) 
hirta var. amboinensis (v.A.v.R.— Gleichenia amboinensis) 26:269 
hirta var. amoena (v.A.v.R.— Gleichenia amoena) 26:269 

. hirta var. candida (Rosénst.—Gleichenia candida) 26:269 


cleFehefelutestutcielctencleltlolutonstolesol relate: 


AAAAAAAAA 


Fern names of R. E. Holttum 227 


(Gleichenia cont.) 


. hirta var. lanuginosa (v.A.v.R.—G. ornamentalis var.) 28:27 

. hirta var. ornamentalis (Rosenst. — Gleichenia ornamentalis) 26:269 

. hirta var, virescens (Hieron. — Gleichenia dolosa var.) 26:270 

. hooglandii 41:327. - NE New Guinea, Hoogland & Schodde 7692 (CANB) 
loheri var. major 26:272.— Celebes, Bunnemeijer 11965 (BO; K, L) 
matthewii 26:265.— Sumatra, Matthew s.n., 31 Jan. 1912 (K) 

microphylla var. vulcanica (Bl. — Gleichenia vulcanica) 24:66 

paleacea (Copel. — Hicriopteris) 26:265 

papuana 26:266.——N. Guinea, Brass 22597 (BM) 

papuana var. membranacea 26:266.— N. Guinea, Brass 24763 (BM) 
peltophora var. schizolepis C. Chr. ex. 26:262.— Sarawak, Mjoberg 9 (P) 
pulchra (Copel. — Sticherus) 26:271 

. Sumatrana 26:264.— Sumatra, Alston [‘1484” sphalm.] 14841 (BM) 
truncata var. bracteata (v.A.v.R.—G. laevigata var.) 26:271 

truncata var. celebica 28:22. — (Central) Celebes, 2600-3000 m. alt. [nothing 
else quoted] 

truncata var. involuta 26:272.— Sumatra, Winkler (Rosenst., Fil. Sumatr. 
exsicc. 182) (L; BM, P) 

truncata var. plumiformis (Presl — Mertensia) 26:272 

. venosa (Copel. — Sticherus) 29:20 

. volubilis var. peninsularis 26:265.— Malay Peninsula, Bintang FMS. Mus. s.n., 
June 1917 (K) 


Grammitis crispatula 16:268.— Malaya, Burkill SFN 12705 (SING) 


Haplodictyum bulusanicum 56:61.— Philippine Is., Luzon, Elmer 16585 (K; BO, 
FL-G, L) 


Heterogonium alderwereltii nom. nov. 18:163.f.1.— Pleocnemia stenosemioides 
. giganteum (Blume — Aspidium) 18:166 

. gurupahense (C. Chr. — Dryopteris sagenoides var.) 25:272 
. pinnatum (Copel. — Stenosemia) 18:163 

sagenoides (Mett. — Aspidium) 18:161.f.2 A&B. 

. Saxicolum (Blume — Aspidium) 18:165. f.2 D. 

. subaequale (Rosenst.— Aspidium) 28:274 

subglabrum 25:273. — Sumatra, Lérzing 5520 (BO) 

. subsageniaceum (Christ — Aspidium) 25:273 

. teysmannianum (Bak. — Acrostichum) Posthumus ex. 19:29 
. wigmanil (Racib. — Aspidium) 25:27] 


Histiopteris caudata (Copel. — Lepidocaulon) 42:459 
H. pilosa 42:460.— Sumatra, Matthew s.n. (K) 
H. squamulata 42:459.— New Guinea, Millar & Holttum 18640 (K) 


Hymenophyllum johorense 3:408 cum fig. — Malaya, Holttum 10755 (SING) 
Lastreopsis parishii var. chupengensis (Ridl. — Lastrea chupengensis) 24:498 
Lepisorus longifolius (Bl.— Grammitis) 24:151 

Leptochilus simplicifolius 3:409.— Malaya, Henderson 17977 (SING) 


Lindsaya cultrata var. parvula 4:61 cum fig. — Malaya, Nur 7297 (SING) 

L. cultrata var. plumula (Ridley. — Lindsaya) 24:329 

L. integra 4:67 cum fig. — Malaya, Holttum 20934 (SING) 

L. kinabaluensis 7:237.— Borneo, Clemens 27744 (simply pinnate), Holttum 
25433 (bipinnate) (SING) 

L. malayensis 4:69 cum fig. — Malaya, Haniff 4032 (SING) 


AAA A AgAgAAAAANDDADAD 


sofaofesfeofasfastesf=sfecfe= 


228 Gardens’ Bulletin, Singapore — XXX (1977) 


(Lindsaya cont.) 


L. scandens var. terrestris 24:327. — (sine typus) 
L. tenera var. gigantea (Hook. — L. flabellata var.) 4:65. 


Lomagramma brassii 38:224.— Papua, Brass 24947 (A) 

L. copelandii 11:201. tt. 9 & 10.— Philippine Is., Ramos & Edafio PBS 28826 
(PNH; BO) 

. cordipinna 11:202. tt. — Samoa, Safford s.n., March 1888 (US) 

cultrata (Baker — Acrostichum) 11:203 

grosseserrata 11:203. tt. 10 & 11.— Siam, Winit 1019 (SING) 

leucolepis 38:224.— W. New Guinea, Brass 12950 (L; BM, BO, MICH, GH) 

. matthewii (Ching — Campium) 11:206 

. merrillii 11:208.t. 14. — Philippine Is., Merrill 8282 (PNH) 

. tahitensis 37:223. — Tahiti, Vesco s.n. (P; B, K) 


omariopsis amydrophlebia (Slosson— Stenochlaena) 14:617 

boivinii 13:53. — Madagascar, Boivin 1947/2 (P) 

congoénsis 14:622. ff. 8 & 9.— Portuguese Congo, Gossweiler 8219 (BM) 
. crassifolia 13:57. Madagascar, Humblot 529 (P; K) 

intermedia (Copeland — Stenochlaena) 6:270 

jamaicensis (Underw. — Stenochlaena) 14:614 

kingii (Copeland — Stenochlaena) 6:273 

kingii var. petiolulata Ros. ex. 6:273.—- New Guinea, Bamler 50 (UC) 
. kunzeana (Underw. — Stenochlaena) 14:617 

lanceolata 14:619. — Brazil, Sellow s.n. (B) 

. latiuscula (Maxon — Stenochlaena) 14:617 

lineata (Presl — Olfersia) 47:9 

longicaudata (Bonap. — Stenochlaena) 13:58 

maxonii (Underw. — Stenochlaena) 14:617 

mexicana 14:617.— Mexico, Hahn 56 (P) 

muriculata 14:624. f.10.— Belgian Congo, Jespersen 26 (BR) 
nigrescens 14:627. ff. 13 & 14.— Fernando Po, Mann s.n. (K) 
nigro-paleata 14:618. ff.4&5.— Peru, Klug 2082 (US; BM, K, NY) 
. raciborskii (C. Chr.) 6:272.— Stenochlaena Raciborskii 

. rossii 14:625. ff. 11 & 12. — Nigeria, Ross 22 (BM) 

. setchellii (Maxon) 6:276. — Stenochlaena Setchellii 

. speciosa 14:620. — Brazil, Sellow 370 (B) 

. Subtrifoliata (Copeland — Stenochlaena) 6:274 

. underwoodii 14:614. ff, 1 & 2.— Jamaica, Jenman s.n. (NY) 


Macrothelypteris ogasawarensis (Nakai — Dryopteris) 48:27 
M. polypodioides (Hook. — Alsophila) 48:29 
M. torresiana var. calvata (Bak. — Nephrodium setigerum var.) 48:28 


Se al al we we ae 


Seepage wieilelgic laluigl ls) |< 


Mesoneuron brackenridgei (Mett.— Aspidium) 44:44 
M. wantotensis (Copel. — Dryopteris) 34:134 


MESOPHLEBION nom. gen. nov. 50:29. — Mesoneuron 

. subgen, PLESIONEURON 50:30. — type species: M. tuberculatum 
. arenicola 63:228.— Sumatra, Meijer 4478 (L) 

. auriculiferum (v.A.v.R.— Dryopteris) 63:231 

. beccarianum (Cesati— Nephrodium) 63:230 

. Carolii 63:227. — Sarawak, Hose 712 (K; BM) 

. chlamydophorum (Rosenst, ex. C. Chr.— Dryopteris) 63:231 

. crassifolium (Bl.) 50:30.— Aspidium crassifolium 

. dulitense 63:229.— Sarawak, Hose s.n., 1894 (K) 

. echinatum (Mett.— Aspidium) 63:226 


—<SS<SS52588 


a 


Fern names of R. E. Holttum 229 


(Mesophlebion cont.) 


. endertii (C. Chr. — Dryopteris) 63:228 

. hallieri (Christ — Aspidium) 63:227 

. motleyanum (Hook. ex. Hook. & Bak.— Nephrodium) 62:209 
. Oligodictyon (Baker — Acrostichum) 63:227 

pallescens (Brause — Dryopteris) 63:230 

. persquamiferum (v.A.v.R.— Dryopteris) 63:226 

. rufescens 63:231.— Papua, Brass 27957 (K; BO, L, LAE) 
teuscheri (v.A.v.R. — Dryopteris) 63:228 

. trichopodum (C. Chr. — Dryopteris) 63:226 

. tuberculatum (Cesati— Nephrodium) 50:30 

. vinosicarpum (v.A.v.R.— Dryopteris) 63:229 


Metathelypteris dayii (Bedd. — Nephrodium) 62:205 
M. fragilis (Baker — Polypodium) 57:127 


Microsorium congregatifolium (v.A.v.R.— Pleopeltis) 24:178 
M. sarawakense (Bak.— Polypodium) 24:175 


NANNOTHELYPTERIS gen. nov. 50:38.— type species: N. aoristisora 

N. aoristisora (Harr. — Polypodium) 50:38 

N. inaequilobata 56:67. — Philippine Is., Luzon, Cuming 272 (K; B, E, G, L, US) 
N. nervosa (Fée — Phegopteris) 56:66 

N. philippina (Presl — Physematium) 56:66 


Oreopteris elwesii (Bak. ex Hook. & Bak. — Nephrodium) 62:204 
Orthiopteris kingii (Bedd. — Dicksonia) 24:306 
Parathelypteris salazica 57:136.— Réunion, Cadet 1981-bis (P) 


PARATHYRIUM gen. nov. 27:449. — species typica: P. boryanum [= Dryoathy- 
rium Ching] 

P. boryanum (Willd. — Aspidium) 27:449 

. forsythii-majoris (C. Chr. — Dryopteris) 27:449 

. macdonellii (Bedd. — Asplenium) 27:449 

. parvisorum (C. Chr.— Dryopteris) 27:449 

. pterorachis (Christ — Athyrium) 27:449 

. unifurcatum (Bak. — Nephrodium) 27:449 

. viridifrons (Makino — Athyrium) 27:449 


Phymatodes crustacea (Copel. —- Lecanopteris) 24:190 
P. subfasciata 16:267.— Malaya, Holttum SFN 21559 (SING) 


Pleocnemia andaiensis (Baker — Polypodium) 61:350 

. brongniartii (Bory — Polypodium) 61:350 

. chrysotricha (Baker — Nephrodium) 20: 187 

. conjugata var. elatior 20:179. — Sumatra, Matthew 512 (BO) 

. dimidiolobata 20: 184. ff. 16, 17. — Solomon Is., Brass 2648 (BO) 

. elegans (Copel. — Tectaria) 61:348 

. hemiteliiformis (Racib. — Pleocnemia leuzeana var.) 20:179. f. 11. 
intermedia 61:349. — Sarawak, Anderson S 32555 (K) 

. irregularis (Presl— Polypodium) 61:347 

macrodonta (Fée — Dictyopteris) 61:348 

. megaphylla 61:357. - New Guinea, Jermy 8149 B (cult. K) 

olivacea (Copel. — Tectaria) 20:181. ff. 8, 10 

. pleiotricha 20: 182. ff. 12, 13. North Borneo, Elmer 21421 (SING; BO) 
. presliana 20: 183. ff. 14, 15. — Philippine Is., Edafio BS 78717 (SING; BO) 
. seranensis 20:187.— Moluccas, Rutten 1850 (BO; SING) 

. submembranacea (Hayata — Aspidium) 61:352 


SSSSS555888 


‘oro ro tS ee 


a-ha-a-Na-la-la-la-Ba-Ba-Ba-Ba-Ba-he-Re-he-) 


230 Gardens’ Bulletin, Singapore — XXX (1977) 


(Pleocnemia cont.) 


E. 
:. 


tripinnata 20: 185, ff. 18, 19. — New Guinea, Lauterbach 560 (SING; BO) 
winitii 20: 181. — Thailand, Winit 36 (SING) 


PLESIONEURON (Holtt.— Mesophlebion subgen. Plesioneuron) 63:232 
P. altum (Brause — Dryopteris) 63:241 


ys yo oO 


POI UVUIUUUUUUUU UU UU UU UU UU VUDUUUVUUUUUUUUUUT 


. archboldiae (Copel. — Lastrea) 63:242 

. attenuatum (Brack. — Lastrea) 63:245 

. belense (Copel. — Dryopteris) 63:239 

. bipinnatum (Copel. — Dryopteris) 63:242 
. costulisorum (Copel. — Lastrea) 63:246 


crassum (Copel. — Dryopteris) 63:243 
ctenolobum 63:247.— NE New Guinea, Jermy 3604 (BM) 


. doctersii 63:246.— W. New Guinea, Docters van Leeuwen 10357 (BO; L) 


dryas 63:247.— Papua, Brass 23447 (BM; L, LAE) 


. dryopteroideum (Brause — Alsophila) 63:237 


falcatipinnulum (Copel. — Dryopteris) 63:244 


. fuchsii 63:236.— N. Borneo, Fuchs 21477 (L; K) 
. fulgens (Brause — Dryopteris) 63:238 
. hopeanum (Bak. — Nephrodium) 63:244 


hopeanum var. acutilobum 63:245.— Fiji, Milne 190 (K) 


. Irayense (Copel. — Cyclosorus) 63:239 


kundipense 63:247. — NE New Guinea, Millar & Holttum NGF 18593 (LAE; 
BRI, K) 


. marattioides (Alston — Dryopteris) 63:238 
. myriosorum (Copel. — Dryopteris) 63:244 


notabile (Brause — Dryopteris) 63:237 


. ophiura (Copel. — Dryopteris) 63:248 
. phanerophlebium (Bak. — Nephrodium) 63:243 
. platylobum 63:246.— NE New Guinea, Jermy 3610 (BM) 


ponapeanum (Hosokawa — Phegopteris) 63:241 
prenticei (Carr. — Lastrea) 63:244 


. pullei 63:236.— W. New Guinea, Pulle 905 (L; BM) 


quadriquetrum (v.A.v.R. — Dryopteris) 63:242 


. rigidilobum 63:248.— NE New Guinea, Walker 774i (BM) 
. royenii 63:240 — W. New Guinea, van Royen 5469 (L) 
. savaiense (Bak. — Nephrodium) 63:240 


septempedale (Alston — Dryopteris) 63:238 
subglabrum 63:243.— Solomon Is., Braithwaite RSS 4512 (K) 
subterminale 63:241. -NE New Guinea, Brass 32049 (K; L, LAE) 


. tahitense 63:241. — Tahiti, Savatier 752 (K) 

. translucens 63:240.— Amboina, Brooks 17760 (BO; BM) 
. tuberculatum (Cesati— Nephrodium) 63:246 

. tuberculatum (Cesati— Nephrodium) 63:246 


wantotense (Copel. — Dryopteris) 63:245 


. wariense (Copel. — Dryopteris) 63:239 
. woodlarkense (Copel. — Cyathea) 63:236 


n 
. angusticaudata 55:308.— Papua, Brass 24984 (BM; LAE) 

. auctipinna 55:310.—N. Celebes, P. & F. Sarasin 119 (BAS) 
. basicurtata 55:309. — Sumatra, Robinson & Kloss 148 (BM) 
. blastophora (Alston — Cyclosorus) 57:156 

. brooksii (Copel. — Dryopteris brooksii) 55:308 

. bryanii (C. Chr. — Dryopteris) 55:317 

. caudata (Holtt.— Pseudocyclosorus) 55:321 


eumatopteris afra (Chr. — Dryopteris) 55:306 


Fern names of R. E. Holttum 231 


(Pneumatopteris cont.) 


. cheesmaniae 55:307.—-W. New Guinea, Cheesman 1269 (BM) 

. christelloides 55:311.— N. Borneo, Mt Kinabalu, Clemens 27541 (K) 

. comorensis 55:318.— Comoro ls., Hildebrandt 1782 (K; B, BM, FI, L) 
. costata (Brack. — Goniopteris) 55:305 

. deficiens 55:321. — Flores, Posthumus 3183 (BO; K) 

dicranogramma (v.A.v.R.— Dryopteris) 55:30] 

ecallosa (Holtt. —- Cyclosorus) 55:310 

egenolfioides 55:306.— W. New Guinea, Brass 11497 (L; BM, BO) 

. excisa (Holtt.— Pseudocyclosorus) 55:32] 

. glaberrima (Richard — Aspidium) 55:318 

. glabra (Copel. — Cyclosorus) 55:301 

glandulifera (Brack. — Goniopteris) 55:302 

hudsoniana (Brack. — Nephrodium) 55:317 

. humbertii 55:313. — Madagascar, Humbert 31679 (P) 

imbricata 55:322.— Amboina, Brooks 17775 (BM; BO, L) 

. incisa 55:317. — Ternate, Alston 16623 (BM) 

. inclusa (Copel. — Dryopteris) 55:306 

. jermyi 55:310.— NE New Guinea, Jermy 3739 (BM) 

kerintjiensis 55:312.— Sumatra, Alston 14149 (BM) 

keysseriana (Rosenst.— Dryopteris) 55:320 

laevis (Mett. — Aspidium) 55:308 

. laticuneata 55:312. - NE New Guinea, Jermy 8044 (BM) 

. latisquamata 55:305.— NE New Guinea, Millar & Holttum NGF 15777 (K: 
LAE) 

. ligulata (Presl — Lastrea) 55:320 

. longipes (Bl. — Aspidium) 55:306 

. lucida (Bak. — Nephrodium) 55:307 

. Macroptera (Copel. — Dryopteris) 55:316 

. magnifica (Copel. — Dryopteris) 55:319 

. mesocarpa (Copel. — Dryopteris) 55:303 

. michaelis 55:313.— Philippine Is., Price 317 (K) 

. Microauriculata 55:311.— N. Borneo, Mt Kinabalu, Clemens 27137 (BM) 
. microloncha (Chr. — Dryopteris) 55:308 

micropaleata 55:319.— N. Borneo, Mt Kinabalu, Holttum 58 (K; SING) 
. nephrolepioides (C. Chr. — Dryopteris) 55:320 

nitidula (Pres] — Nephrodium) 55:318 

novae-caledoniae 55:313. — New Caledonia, Lécard s.n., Oct. 1876 (P) 
obliqua 55:309. — Celebes, P. & F. Sarasin 127 (BAS) 

obstructa (Copel. — Dryopteris) 55:302 

. oppositifolia (Hook. — Polypodium) 55:304 

. oxyura (Copel. — Dryopteris) 55:315 

petrophila (Copel. — Dryopteris) 55:322 

. regis (Copel. — Dryopteris) 55:322 

. remotipinna (Bonap. — Dryopteris) 55:311 

. rodigasiana (T. Moore — Nephrodium) 55:319 

rotumaensis (St. John — Cyclosorus) 55:318 

papuana 55:311.— Papua, Brass 22778 (LAE; BM, L) 

. parksii (Ballard — Dryopteris) 55:302 

. patentipinna 55:307.— NE New Guinea, Walker 8477 (BM) 

. pennigera (Forst.— Polypodium) 55:305 

. pergamacea 55:315.— New Ireland, Jermy 7877 (cult. K, 123/70, no. 1035) 
. prismatica (Desv.— Nephrodium) 55:316 


OOOO UOMO UU UU UU UUM VUVUUVUUUUMUUUUUUUUU UU UY 


232 Gardens’ Bulletin, Singapore — XXX (1977) 


(Pneumatopteris cont.) 


a-Ma-Ba-Ma-Na-Ba-Be-Ba- Bene Be- eee 


a-Na-Ra-Me-Me- ha" 


. sandwicensis (Brack. — Stegnogramma) 55:303 

. sibelana 55:301.— Moluccas, Batjan, Alston 16942 (BM) 
. sogerensis (Gepp. — Dryopteris) 55:315 

. stokesii (E. Brown — Dryopteris) 55:302 

. subappendiculata (Copel. — Dryopteris) 55:303 

. subpennigera (C. Chr. — Dryopteris) 55:304 

. sumbawensis (C. Chr. — Dryopteris) 55:323 

. Superba (Brause — Dryopteris) 55:303 

. remotipinna (Bonap. — Dryopteris) 57: 159 

. tobaica 55:316. — NE Sumatra, Surbeck 14 (L; BO) 
. transversaria (Brack. — Nephrodium) 55:319 

. truncata (Poir. — Polypodium) 55:314 


truncata var, loyalii 55:314. — Darjeeling, Loyal sn. Aug. 1956 (Herb. Punjab 
Univ. no. 3172) 


unita (Kunze — Gymnogramma) 55:304 


. usambarensis 55:312. — Tanzania, Faden et al. 70/293 (EA) 

. vaupelii (C. Chr. — Dryopteris) 55:317 

. venulosa (O, Kuntze — Dryopteris) 55:315 

. versteeghii 55:321. — W. New Guinea, Versteegh BW 10259 (L) 
. walkeri 55:323. — NE New Guinea. Walker 9980 (BM) 


PODOSORUS gen. nov. 42:455.— Type-species: P. angustatus Holtt. 


P. 


angustatus 42:455. — Philippine Is., Gutierrez PNH 78332 (K; L) 


Polypodium papilliferum 2:22.— Mentawi Is., Kloss 14800 (SING) 


POLYSTICHOPSIS (J. Sm. pro subsect. sub Lastrea) — 17:149, 152 [= Arachnio- 


SO oO *O Fo 


des BI.] 


. aristata (Forst. — Polypodium) 24:468 

. chinensis (Rosenst. — Polystichum amabile var.) 24:486 
. hanifii (Holtt.— Dryopteris) 24:485 

. hasseltii (Bl. — Polypodium) 24:487 

. puncticulata (v.A.v.R. — Polystichum) 24:487 


Pronephrium sect. DIMORPHOPTERIS (Tagawa & K. Iwats. — Dimorphopteris 


ee eS OO eo OU 


pro gen.) 50:36. ff. 10, 10a 


. sect, GRYPOTHRIX 50:36. fig. 8. — type species: Meniscium cuspidatum BI. 
. acanthocarpum (Copel. — Dryopteris) 53: 107 

. amboinense (Willd. — Aspidium) 53:120 

. aquatiloides (Copel. — Dryopteris) 53:108 

. articulatum (Houlst. & Moore — Nephrodium) 53:116 

. asperum (Presl — Polypodium) 53:112 

. brauseanum nom, nov. 53:107.— Dryopteris canescens var. novoguineensis 
. beccarianum (Cesati— Meniscium) 53:107 

. buwaldae 53:115.— Aru Is., Buwalda 4979 (L) 

. celebicum (Bak. — Acrostichum) 53:119 

. clemensiae (Copel. — Dryopteris 53:118 

. crenulatum 53:123.— Vietnam, Eberhardt s.n. 1896 (P) 

. cuspidatum (Bl. — Meniscium) 53:123 

. debile (Bak. — Nephrodium) 53:118 

. diminutum (Copel. — Dryopteris) 53:115 

. elmerorum (Copel. — Dryopteris) 53:118 


Fern names of R. E. Holttum 233 


(Pronephrium cont.) 


DS WUD UUD UD UND UN DUD DU UU UU UD 


. euryphyllum (Rosenst. — Dryopteris) 53:112 

. exsculptum (Bak. — Acrostichum) 53:117 

. firmulum (Bak. — Polypodium) 53:116 

. gardneri 49:81.— Ceylon, Gardner 1135 (K), s.n. (BM) 


glandulosum (Bl. — Aspidium) 53:118 
granulosum (Presl — Polypodium 53:119 
gymnopteridifrons (Hayata — Dryopteris) 53:112 
hosei (Bak. — Meniscium) 53:120 


. insulare (K. Iwats. — Abacopteris) 53:123 


kjellbergii 53:118.— Celebes. Kjellberg 2638 (BO) 
lakhimpurense (Rosenst. — Dryopteris) 53:110 
longipetiolatum (K. Iwats.— Abacopteris) 53:123 
megacuspe (Bak. — Polypodium) 53:122 
melanophlebium (Copel. — Dryopteris) 53:108 
menisciicarpon (Bl. — Aspidium) 53:111 


. merrillii (Chr. — Dryopteris) 53:117 


micropinnatum 53:108.— NE New Guinea, Darbyshire & Hoogland 8014 
(CANB; BM, L, LAE) 

millarae 53:115.— NE New Guinea, Womersley & Millar NGF 8500 (LAE) 

minahassae 53:119.—N. Celebes, Alston 16570 (BM) 

moniliforme (Tagawa & K. [wats.— Dimorphopteris) 50:36 


. nitidum (Holtt. — Dryopteris urophylla var.) 53:109 


nothofageti 53:118.— NE New Guinea, Walker 8788 (BM) 
nudatum (Roxb. — Polypodium 53:111 

palauense (Hosok. — Meniscium) 53:121 

palopense 53:116.— Celebes, Kjellberg 1999 (BO) 

parishii (Bedd. — Meniscium) 53:12 

penangianum (Hook. — Polypodium) 53:110 

pentaphyllum (Rosenst. — Dryopteris) 53:108 

peramalense 53:115.— W. New Guinea, Pulle 415 (BM: L) 
repandum (Fée — Goniopteris) 53:109 

rhombeum (Chr. — Dryopteris diversifolia var. acrostichoides subvar. 53:120 
rubicundum (v.A.v.R. — Phegopteris) 53:123 

rubidum (J. Sm. ex Hook. — Polypodium) 53:111 

rubrinerve (Mett. — Phegopteris) 53:110 


. Salicifolium (Wall. ex Hook. — Meniscium) 53:123 


samarense (Copel. — Cyclosorus) 53:119 
simillimum (C. Chr. — Dryopteris) 53:116 


. simplex (Hook. — Meniscium) 53:122 

. solsonicum 64:197.— Philippine Is., Luzon, Price 2903 (K; PNH) 

. tibangense (C. Chr. — Dryopteris) 53:118 

. thwaitesii (Hook. — Meniscium) 53:122 

. triphyllum (Sw. — Meniscium) 53:122 

. womersleyi 53: 108. NE New Guinea, Floyd & Womersley NGF 6308 (LAE: 


BM) 


. xiphioides (Chr. — Dryopteris) 53:119 


Pseudocyclosorus camerounensis 57:137.— Cameroun, Meurillon CNAC 726 


vUuWUUU 


(P; K 


. caudatus 34: 133. fig. 7.— New Guinea, Holttum, cult. Kew 545/63, no. 12 (K) 
. excisus 34: 133. — New Guinea, Millar & Holttum 18623 (K) 

. johannae 57:138.— Comoro Is., Johanna Id., Bewsher 24 (K) 

. ochthodes (Kunze — Aspidium) 62:204 

. petrophila (Copel. — Dryopteris) 34:133 

. pulcher (Bory ex Willd. — Aspidium) 57: 138 


234 Gardens’ Bulletin, Singapore — XXX (1977) 


Pseudophegopteris aubertii (Desv.— Polypodium) 48:18 
P. cruciata (Willd. — Aspidium) 48:21 
. cyclocarpa 34:131. fig. 10.— New Guinea, Holttum, cult. Kew 578/63, no. 57 
K 


P 
(K) 

P. dianae (Hook. — Polypodium) 48:21] 

P. henriquesii (Bak. — Polypodium) 48:15 

P. hirtirachis (C, Chr. — Dryopteris) 48:22 

P. keraudreniana (Gaud.— Polypodium) 48:19 

P. kinabaluensis 48:16.— Borneo, Fuchs 21475 (KSEPL; K, L) 

P. persimilis (Bak. — Polypodium) 48:20 

P. procera (Mann — Phegopteris) 48:18 

P. pyrrhorhachis var. glabrata (Clarke — Polypodium distans var.) 48:24 

P. rectangularis (Zoll. — Polypodium) 48:19 

P. sumatrana 48:22. — Sumatra, Holttum 26211 (K; SING, BO) 

P. tenggerensis 58:500.— Java, Buysmann 40 (U) 


Pteris simplex 31:531.— E. New Guinea, Cruttwell 985 (K) 
P. werneri (Rosenst. — Hemipteris) 42:457 


Pyrrosia costata (Presl — Apalophlebia costata 1851) 47:27 

[transfer previously made in 1967, with Niphobolus costatus [Presl] Pres] ex Bedd. 
(1866) as basionym, by Tagawa & Iwatsuki in Act. Phytotax. Geobot. 
22: 100] 

P. penangiana (Hook. — Niphobolus) 24: 146 

P. stictica (Kze — Niphobolus) 47:31 


Rheopteris asplenioides 31:531. -SW New Guinea, Kalkman BW 8528 (L) 
Rumohra puncticulata (v.A.v.R. — Polystichum) 9:137 
Schizaea paucijuga 16:267. — Malaya, Haniff SFN 1098 (SING) 


Schizoloma javanense (Bl. — Lindsaya) 24:349 
S. tenerum (Dryander — Lindsaya) 24:348 


Selliguea lima (v.A.v.R. — Pleopeltis) 24:157 


Sphaerostephanos angustifolius (Pres]l— Nephrodium) 65:66 
. appendiculatus (Blume — Gymnogramma) 65:62 

. arbuscula (Willd. — Aspidium) 57: 164 

. dichrotrichoides (v.A.v.R. — Dryopteris) 65:65 

. diversilobus (Presl — Nephrodium) 65:57 

. elatus (Bojer — Aspidium) 57: 167 

. elatus var. thomensis 57: 167. — Sao Tomé, Moller s.n. 1885 (P) 
. ellipticus (Rosenst. — Dryopteris) 65:66 

. elmerorum (Copel. — Dryopteris) 65:55 

. erectus (Copel. — Cyclosorus) 65:64 

. fenixii 65:55. — Batanes Is., Fenix BS 3780 (P; US) 

. heterocarpus (Blume — Aspidium) 62:209 

. heterocarpus var. bordenii (Christ — Dryopteris) 65:67 

. hirsutus (Kunze ex Mett.— Aspidium) 65:63 

. lastreoides (Pres] — Pronephrium) 65:54 

. latebrosus (Kunze ex Mett.— Aspidium) 62:209 

. lobatus (Copel. — Cyclosorus) 65:64 

. loherianus (Christ — Aspidium) 65:61 

. penniger (Hook. — Nephrodium) 62:209 

. magnus (Copel. — Cyclosorus) 65:61 

. mindorensis 65:55. — Philippine Is., Mindoro, Price 690 (K; PNH) 
. perglanduliferus (v.A.v.R. — Dryopteris) 65:59 


NANNNNNANNUNUAUNNURNUNUURAURNRNURNRUNUURNN 


Fern names of R. E. Holttum 235 


(Sphaerostephanos cont.) 


. polisianus 65:65. — Philippine Is., Luzon, Price 431 (K; PNH) 

. polycarpus (Blume — Aspidium) 65:56 

. porphyricola (Copel. — Dryopteris) 65:59 

. productus (Kaulf.— Aspidium) 65:59 

. punctatus 65:60.— Philippine Is., Luzon, Elmer 16642 (US; BO, G, K, L, 
SYD) 

. Sakayensis (Zeiller — Nephrodium) 62:209 

. santomasii 65:62. — Philippine Is., Luzon, Price 1034 (K; PNH) 

. sessilipinna (Copel. — Dryopteris) 65:53 

. simplicifolius (J. Sm. ex Hook. — Aspidium) 65:61 

. smithianus (Pres! — Nephrodium) 65:65 

. spenceri (Christ — Dryopteris) 65:58 

. stenodontus (Copel. — Cyclosorus) 65:56 

. subtruncatus (Bory — Polypodium) 48:80 

. tephrophyllus (Copel. — Dryopteris) 65:58 

. todayensis (Christ — Dryopteris) 65:58 

. unitus (L. — Polypodium) 57:165 

. williamsii (Copel. — Dryopteris) 65:53 


Stenochlaena cumingii 51:122.— Philippines Is., Cuming 226 (K; BM) 
Syngramma minima 1:56.— Malaya, Holttum 17498 (SING) 


Taenitis cordata (Gaud. — Schizoloma) 29: 112 

T. dimorpha 16:274. — Malaya, Holttum SFN 17399 (SING) 

T. diversifolia 46:93. — Solomon Is., RSS 4360 (K) 

flabellivenia (Bak. — Polypodium) 31:533 

. hosei (Bak. — Hemionitis) 46:92 

lanceolata (Diels — Syngramma) 46:91 

marginata 31:532.— Sumatra, Ernst 701 (Z; K) 

pinnata (J. Sm.— Syngramma) 46:90 

pinnata var, brachysora (Bak. —- Gymnogramme) 46:91 

pinnata var. polypodioides (Bak.— Gymnogramme) 46:91. fig. 4 

trilobata nom. nov. 46:93. — Lindsaya trilobata Bak. non Colenso (= L. hosei 
C. Chr. non Taenitis hosei (Bak.) Holtt.) 


vittarioides 42:460. — Sumatra, Korthals (L; K) 


Tectaria grandidentata (Cesati — Polypodium dilatatum var.) 24:514 

T. gymnosora 43:308. — Thailand, Hansen, Seidenfaden & Smitinand 11225 (C; K) 
T. ingens (Atkinson ex Clarke — Nephrodium) 24:503 

T. isomorpha 44:49. — Bismarck Archipelago, Noona Dan Exped. 2369 (C) 


TERATOPHYLLUM Mett. emend. 12:355.— Type species: Lomaria aculeata 


NANNNNNNNANHNNM ANNNNMN 


5 


T. aculeatum var. montanum 6:289, text. ff. 34, 41; tt. 1 & 4.— Malay Peninsula, 
Kunstler 8309 (SING) 

T. arthropteroides (Christ — Stenochlaena) 6:303. t. 12 

T. brightiae (F. von Muell. — Acrostichum) 12:358. t. 28. 

T. clemensiae 7:262, text-ff, 1-9. — Borneo, Clemens 31614 (SING) 

T. gracile (Blume — Lomaria) 6:291. text-ff. 4, 42 t. 5. 

T. koordersii 6:301. text-ff. 48 & 49. t. 11.—Celebes, Koorders 17065B (adult 
fronds), 17064B and 17123B (bathyphylls) (BO) 

T. leptocarpum (Fée — Lomariopsis) 10:143 

T. ludens (Fée — Lomariopsis) 6:298. text-ff, 5, 23-28, 31, 35, 46, 47; t. 10 

T. luzonicum 6:297. t. 9. — Philippine Is., Fenix, Bur. Sci 28272 (BO) 

T. rotundifoliatum (R. Bonap. — Stenochlaena) 6:294. text-ff. 36, 37, 44. tt. 7 & 8. 


236 Gardens’ Bulletin, Singapore — XXX (1977) 


(Teratophyllum cont.) 


T. wilkesianum (Brack. — Polybotrya) 12:359. tt. 29 & 30 
T. williamsii (Underwood — Stenochlaena) 6:292, text-f. 43. t. 6 


Thelypteris herbacea 16:268.— Malaya; Holttum SFN 20571 (SING) 

. khamptorum 49:82.— Burma, Toppin 4217 (K) 

novae-hiberniae 44:50. — Bismarck Archipelago, Noona Dan Exped. 1848 (C) 
. paleata (Copel. — Dryopteris) 24:249 

. motleyana (Hook. — Nephrodium) 24:247 

. unidentata (Bedd. — Lastrea) 24:25] 


TRIGONOSPORA gen. nov, 50:29. ff. 4, 4a. — type species: T. ciliata 
T. calcarata (Blume — Aspidium) 59:506 

T. ciliata (Benth. — Aspidium) 50:29 

T. koordersii (Christ — Aspidium) 59:506 

T. sericea (Scott ex Bedd. — Lastrea) 62:204 


Vaginularia_acrocarpa 44:51.— Bismarck Archipelago, Noona Dan Exped. 
2452 AMS) 


Vittaria angustissima 16:274.— Malaya, Carr s.n., March 1929 (SING) 
V. malayensis 3:409 cum fig. —- Malaya, Burkill & Holttum 8705 (SING) 


Xiphopteris hieronymusii (C. Chr. — Polypodium) 24:218 
X. sparsipilosa (Holtt.— Ctenopteris) 24:219 


HH 


Bibliography 
(dates of publication given here may differ from those given on title-pages) 


A new fern from the Malay Peninsula. Gdns’ Bull. S.S. 4: 56-57. (1927). 
Spolia Mentawiensia: Pteridophyta. J. Mal. Br. R. Asiatic Soc. 6: 14-23. (1928). 
New species of ferns from the Malay Peninsula. Gdns’ Bull. S.S. 4: 408-410. (1929). 


The genus Lindsaya in the Malay Peninsula, with descriptions of two new species. 
Gdns’ Bull. S.S. 5: 58-71. (1930). 


5 Aspidium maingayi (Baker) Holttum comb. nov. Gdns’ Bull. S.S. 5: 207-211. (1931). 


6 On Stenochlaena, Lomariopsis and Teratophyllum in the Malayan Region, Gdns’ Bull. 
S.S. 5: 245-313. (1932). 

7 The ferns of Mt. Kinabalu (C. Christensen & R.E.H.). Gdns’ Bull. S.S. 7: 191-324. 
(1934). 

8 The tree-ferns of the Malay Peninsula. Gdns’ Bull. S.S. 8: 293-320. (1935). 

9 Notes on Malayan ferns, with descriptions of 5 new species. Gdns’ Bull. S.S. 9: 119-138. 
(1937). 

10 Further notes on Stenochlaena, Lomariopsis and Teratophyllum. Gdns’ Bull. S.S. 9: 
139-144. (1937). 

11 The genus Lomagramma. Gdns’ Bull. S.S. 9: 190-221. (1937). 

12 A re-definition of the genus Teratophyllum. Gdns’ Bull. S.S. 9: 355-362. (1938). 


13. The genus Lomariopsis in Madagascar and the Mascarene Islands. Notulae Systematicae, 
Paris 8: 48-62. (1939). 


14. New species of Lomariopsis. Kew Bull. 1939: 613-628. (1940). 

15 The fern genus Diplazium in the Malay Peninsula. Gdns’ Bull. S.S. 11: 74-108. (1940). 
16 2 uma of vascular plants from the Malay Peninsula. Gdns’ Bull. Sing. 11: 267-298. 
17 A revised classification of Leptosporangiate ferns. J. Linn Soc. Bot. 53: 123-158. (1947). 
18 The fern-genus Heterogonium Presl. Sarawak Museum J. 5: 156-166. (1949). 

19 Further notes on the fern-genus Heterogonium Presl. Reinwardtia 1: 27-31. (1950). 

20 The fern-genus Pleocnemia Presl. Reinwardtia 1: 171-189. (1951). 


21 The fern-genus Arcypteris Underwood (Dictyopteris Pres] sensu Fée). Reinwardtia 1: 
191-196. (1951). 


22 A new Malayan Athyrium, Kew Bull. 8: 545. (1954). 


hk WwWne 


Fern names of R. E. Holttum 237 


23 
24 


25 


26 


2A 


28 


29 


30 


31 
32 
33 


34 


35 


36 


37 


38 
39 
40 


41 
42 
43 


a4 


45 


46 
47 


48 


49 


50 


51 


52 


53 


54 


A new fern from Malaya (Athyrium angustipinna). Gdns’ Bull. Sing. 14: 8. (1953). 


A revised Flora of Malaya vol. 2. Ferns of Malaya. Singapore: Government Printing 
Office. (1955). 


Some additional species of Heterogonium. Reinwardtia 3: 269-274. (1955). 


Florae Malesianae Praecursores, XVI: On _ the taxonomic subdivision of the 
Gleicheniaceae, with descriptions of new Malaysian species and varieties. Reinwardtia 4: 
257-280. (1957). 


Notes on Malaysian ferns, with descriptions of a new genus and new species. Kew Bull. 
13: 447-455. (1959). 


Flora Malesiana, Ser. II, Pteridophyta, vol. 1, pt. 1: i-xxiii, 1-61. (Introductory Note; 
List of genera of Malayan Pteridophyta; The Morphology of Ferns; General Keys 
1 & 2; Keys to Genera; Bibliography; Gleicheniaceae, Schizaeaseae). (1959). 


Two problem-species: Schizoloma cordatum Gaud. and Syngramma pinnata J. Sm. 
Amer. Fern J. 50: 109-113. (1960). 


New species of tree-ferns (Cyathea Sm. and Dicksonia L’Herit). Kew Bull. 16: 51-64. 
(1962). 


New species of Malaysian ferns. Blumea 11: 529-534. (1963). 
Cyatheaceae. Flora Malesiana, Ser. II, 1, pt. 2: 65-176. (1963). 


The tree-ferns of the genus Cyathea in Australasia and the Pacific. Blumea 12: 241-274. 
(1964), 


Cytological observations on ferns from New Guinea, with descriptions of new species 
(R.E.H. & S. K. Roy). Blumea 13: 129-139. (1965). 


Tree-ferns of the genus Cyathea Sm. in Asia (excluding Malaysia). Kew Bull. 19: 463-487. 
(1965). 


Studies in the Flora of Thailand 33: Filicinae excl. of the Ophioglossaceae (second list). 
Dansk Bot. Ark. 23: 227-244. (1965). 


The genera Lomariopsis, Teratophyllum and Lomagramma in the islands of the Pacific 
and Australia. Blumea 14: 215-223, (1966). 


Two new species of Lomagramma from New Guinea. Blumea 14: 224. (1966). 
The genus Arthropteris J. Sm. in Malesia. Blumea 14: 225-229. (1966). 


Florae Malesianae Precursores XLIV: The fern-genus Elaphoglossum in Malesia, with 
descriptions of new species. Blumea 14: 317-326, (1967). 


Four new species of ferns from New Guinea. Blumea 14: 327-329. (1967). 
New ferns from Malesia. Kew Bull. 20: 455-460. (1967). 


Studies in the Flora of Thailand 38: Some plants new or rare to Thailand with descrip- 
tions of new species: Filicinae. Dansk Bot. Ark. 23: 308-309. (1966). 


Botanical Report of the Danish Noona Dan Expedition, Filicinae. Dansk Bot. Ark. 25: 
35-52. (1967). 


A revised Flora of Malaya vol. 2. Ferns of Malaya, ed. 2. Singapore: Government 
Printing Office. (1968). 


A re-definition of the fern-genus Taenitis Willd. Blumea 16: 87-95. (1968). 


A commentary on some type specimens of ferns in the Herbarium of K. B. Presl. 
Novitates Botanicae Inst. Bot. Univ. Carol. Prag. 1968: 3-57. (1969). 


Studies in the family Thelypteridaceae 1. The genera Phegopteris, Pseudophegopteris 
and Macrothelypteris. Blumea 17: 5-32. (1969). 


New species of Thelypteridaceae from India, Ceylon and Burma (R.E.H, & P. Chandra). 
Kew Bull. 26: 79-82, (1971). 


Studies in the family Thelypteridaceae 3. A new system of genera in the Old World. 
Blumea 19: 17-52. (1971). 


The genus Stenochlaena J. Sm., with description of a new species. American Fern J. 61: 
119-123. (1971). 

Morphology and anatomy of the genera Davallia, Araiostegia and Davallodes, with a 
discussion on their affinities. (T. & U. Sen & R.E.H.). Kew Bull. 27: 217-243. (1972). 


Studies in the family Thelypteridaceae 4. The genus Pronephrium Pres]. Blumea 20: 
105-126 (1972). 


The identity of three type specimens in the Willdenow Herbarium. Amer, Fern J. 63: 
81-84 (1973). 


238 Gardens’ Bulletin, Singapore — XXX (1977) 


55 Studies in the family Thelypteridaceae 5. The genus Pneumatopteris Nakai. Blumea 21: 
293-325. (1974). 


56 Studies in the family Thelypteridaceae 6. [The genera] MHaplodictyum and 
Nannothelypteris, Kalikasan 2: 58-68. (1974). 


57 Thelypteridaceae of Africa and adjacent islands. $. African J. Bot. 40: 123-168. (1974). 
58 Additions to the Fern Flora of Java. Reinwardtia 8: 499-501. (1974). 

59 The genus Trigonospora (Thelypteridaceae) in Malaysia. Reinwardtia 8: 503-507, (1974). 
60 Studies in the family Thelypteridaceae 7. The genus Chingia. Kalikasan 3 : 13-28. (1974). 
61 The fern-genus Pleocnemia. Kew Bull. 29: 341-357. (1974). 


62 [new thelypterid nomenclatural combinations] in B. K. Nayar, A revised nomenclature 
of the thelypteridoid ferns. pp 202-212 in B. K. Nayar & S. Kaur, Companion to 
R. H. Beddome’s Handbook to the ferns of British India, Ceylon and the Malay Peninsula. 
Koenigstein & New Delhi (1974). 


63 Studies in the family Thelypteridaceae 8. The genera Mesophlebion and Plesioneuron. 
Blumea 22: 223-250 (1975). 


64 New Philippine ferns. Kalikasan 3: 196-197 (1975). 


65 Studies in the family Thelypteridaceae 9. The genus Sphaerostephanos in the Philippines. 
Kalikasan 4: 47-68 (1975). 


Philippine Dryopteris 
by 


M. G. PRICE 


Department of Botany 
University of Michigan 
Ann Arbor, Michigan, U. S. A. 


Summary 


A key to the thirteen recognized Philippine species of Dryopteris is presented, with 
descriptions of two new species, D. uropinna and D. permagna, and four species not hitherto 
reported as Philippine: D. chrysocoma, D. pulvinulifera, D. polita, and D. formosana. 
Ctenitis mearnsii is reduced to Nothoperanema hendersonii. 


The genus Dryopteris Adanson (nom. cons.) is here circumscribed in the 
strict sense adopted by most recent authors, with its nearest relatives in Asia being 
along one lineage Arachniodes and on another Nothoperanema. Dryopteris consists 
of about two hundred species, mostly north subtropical and temperate, with the 
centre of diversity the Himalayan-East Asian region. Distinguishing characters of 
the genus are: 


Rhizome short-erect, infrequently short-creeping, stipes with 5 or more 
vascular bundles at base, axes grooved above, junctions between grooves of different 
orders of axes open, lamina pinnate to quadripinnate, architecture usually anadro- 
mous at proximal base, then catadromous upwards and outwards, or infrequently 
anadromous throughout. Hairs not acicular, nor multicellular and erect, margins 
not bearing long sharp cartilaginous teeth, sinuses without cartilaginous membranes 
Or protuberances, veins free, not reaching margin. Sori along veins, round, 
receptacles strongly raised, indusium round-reniform with a short sinus, somewhat 
stalked (rarely much reduced), paraphyses usually present, spores monolete, 
variously winged, bullate, or tuberculate. 


Arachniodes is always rather strongly anadromous throughout, with dimidiate 
pinnules (basiscopic base excised), and usually with sharp cartilaginous teeth. It 
is, however, extremely difficult to formulate a verbal boundary; among Philippine 
plants Arachniodes hasseltii has blunt or rounded segments and Dryopteris pulvinuli- 
fera and D. subarborea are anadromous. 


Nothoperanema has often been immersed in Dryopteris or Ctenitis, It agrees 
with Dryopteris in soral characters but is vegetatively similar to Peranema and 
Diacalpe, and thence to Acrophorus and Stenolepia. The patent stipe paleae, erect 
Diacalpe, and thence to Acrophorous and Stenolepia. The patent stipe paleae, erect 
conical multiseptate hairs on lamina, and fine dissection clearly mark the genus. It 
might more properly be reduced to Peranema (as has Diacalpe) than Dryopteris. 
Since Nothoperanema has not been reported by name from the Philippines, I 
provide below a brief synonymy and description of the one Philippine species: 


Nothoperanema hendersonii (Bedd.) Ching, Acta Phytotax. Sinica 11 (1966) 28. 
Lastrea hendersonii Bedd., Ferns Brit. India Suppl. (1876) 17, t. 377. —— 
Dryopteris hendersonii (Bedd.) C. Chr., Ind. Fil. (1905) 270; Tagawa, Acta 
Phytotax. Geobot. 6 (1937) 155. —— Ctenitis mearnsii Copel., Philip. J. Sci. 81 
(1952) 24; Fern Fl. Philip. 2 (1960) 289. 


239 


240 Gardens’ Bulletin, Singapore — XXX (1977) 


Rhizome ascending, paleae brown, to 18 X 3 mm, concolorous, with minute 
marginal projections. Stipes bright brown, to 70 cm long and 8 mm thick at base, 
paleae linear, patent, brown, subentire, Lamina subquadripinnate, to 70 cm long, 
ovate-deltoid, anadromous in sequence of pinnules at base, upwards catadromous, 
rachis with reduced linear patent paleae, these grading to stout conical erect 
multiseptate hairs forming a tuft inside rachis groove at junctions, also along 
costae, costules, and veins both sides of lamina; texture thin papyraceous. Basal 
pinnae to 30 cm long, stalk 3 cm, apex acuminate; basal basiscopic pinnule to 
9 cm, secondary pinnules very deeply lobed. Middle pinnae ascending, 12-17 cm 
long, acuminate, stalk 5-10 mm, basal pinnules c. 3 X 1.7 cm, secondary pinnules 
adnate, blunt, shallowly lobed or crenate-serrate, apex toothed. Sori inframedial, 
indusium dark brown, minutely farinose, 0.7 mm across. 


LUZON. Mt Pulog, Curran et al. FB 16277 (MICH), Jacobs 7301 (A, CAHUP, 
PNH); Mt Burnay, Iwatsuki et al. P-774; Benguet, Pauai, McGregor BS 8507 
(MICH); Haight’s Place, Mearns BS 4216 (MICH), 4218 (MICH); Mt Singakalsa, 
Celestino & Castro PNH 3971 (MICH); Mt Santo Tomas, Price 1868; Bulasan, 
Copeland s.n. 7 May 1913 (MICH). 


Ecology: Terrestrial in mountain forests, 1900-2700 m. 
Distribution: N. India, Nepal, S. China, Japan, Thailand, Java. 


Most of the Philippine species of Dryopteris are found in the Benguet highlands 
of Luzon; none were collected by botanists until the 1890s. It is likely that 
additional species remain to be discovered in the mountains of northern Luzon. 


Aside from the delimitation of Dryopteris from Arachniodes, the greatest 
difficulty still remaining is the proper treatment of the D. subarborea complex 
(species 11-13 below). There are at least three Philippine taxa but their relationship 
with plants from elsewhere in Malesia and Polynesia remains to be ascertained. I 
am not sure that I have applied the names D. subarborea and D. purpurascens 
correctly. 


Instead of a full synonymy, I have provided all Philippine references, original 
and basionym citations, and reference to important descriptions. Philippine 
specimens I have seen are cited; duplicates of my own collections are being 
distributed, Ecological information is based on Philippine conditions. 


KEY TO THE PHILIPPINE SPECIES OF DRYOPTERIS 


1. Indusium c. 2 mm across, persistently helmet-shaped; rhizome paleae orange- 


colored 
2. Fronds dimorphic (the fertile lamina contracted), deltoid-ovate, not 
glandular 2102221, 234 Se Se 1. D. cochleata 


2. Fronds uniform, lanceolate, bearing small capitate glandular hairs on axes, 
lamina, ATIC? TOSI eh ae Sec 2. D. chrysocoma 
1. Indusium exceptionally to 1.5 mm across, shriveling; paleae not orange 


3. Lamina lanceolate or narrower, pinnate or bipinnatifid; basal pinnae similar 
to middle pinnae 


4. Pinnae toothed or shallowly lobed ........................005 3. D. hirtipes 
4. Pinnae lobed nearly to COSEA, ses.) aniee a) eee 4. D. wallichiana 


3. Lamina ovate or deltoid-ovate, at least bipinnate; basal pinnae more 
dissected than middle pinnae, considerably longer and/or broader 


Philippine Dryopteris 241 


5. Lamina to c, 50 cm long 


6. Lamina anadromous throughout; rhizome short-creeping, paleae 
golden-brown, acicular ........................0.0008 5. D. pulvinulifera 


6. Lamina catadromous above base; rhizome short-erect, paleae brown 
or black 

7. Stalks of basal pinnae over 15 mm long; indusium minute 
len AMR Seer eamledalle: seehpeaee lip eeetet Beas 6. D. polita 

7. Stalks of basal pinnae 10 mm or less long; indusium evident 
8. Paleae of rhizome and stipe pale brown, very thin, lanceo- 
late or ovate; rachis groove strongly papillate within ...... 
ee a oe ae 7. D. sparsa 


8. Paleae at least partially black, not thin, linear or narrowly 
lanceolate; rachis groove not papillate 


9. Basal basiscopic pinnule of basal pinnae about equal 
to succeeding basiscopic pinnules 


9. Basal basiscopic pinnule much enlarged 


10. Pinnules mostly somewhat falcate (costules ascend- 
a tale sea I ahaa 9. D. varia 


10. Pinnules never falcate (costules not arcuate) ...... 
bd hd COS ERS teers, |. 5: 10. D. formosana 


5. Lamina over 100 cm long 
11. Rhizome short-creeping; lamina glabrous 


eres eee e eee eee eee eee er eeeeeeee 


11. Rhizome short-erect; lamina with appressed multi-cellular hairs 


beneath 
12. Sori containing conspicuous paraphyses with large yellow 
EE i oan ans 12. D. purpurascens 


12. Paraphyses inconspicuous, hyaline, shorter than sporangia, 
heads not or only very slightly expanded ......................5. 
baboons: DYE | ate b at rcect eh dy Kocsis iiteeiaice Lo. DD.’ permagna 


1. Dryopteris cochleata (Don) C. Chr., Ind. Fil. (1905) 258; Ching, Bull. Fan 
Mem. Inst. Biol. Bot. 8 (1938) 434; Copel., Fern Fl. Philip. 2 (1960) 282. —— 
Nephrodium cochleatum Don, Prod, Fl. Nepal. (1825) 6. D. heleopteroides 
Christ, Philip. J. Sci. 2C (1907) 212; Merrill & Merritt, Philip. J. Sci. 5C 
(1910) 317. 


LUZON. Benguet, Bacani FB 15962 (MICH), Copeland 1837a (MICH), Hernaez 
CAHUP 13898, Price 1146, 1823, 1827, Ramos BS 5813 (NY, US, MICH), 
Williams 1502 (NY, US); Bontoc, Vanoverbergh 3343 (F); Ilocos Norte, Iwatsuki 
et al. P-113, P-151, P-221, Price 3341. 


Ecology: Rhizome short-creeping, terrestrial or on limestone, 200-1800 m, in fairly 
open seasonally dry grassland, under pine forest, or at borders of secondary growth, 
in sites with good drainage, cool nights, and no wind. 


Notes: Copeland described D. cochleata as glabrous but the lamina beneath has 
appressed multicellular hairs identical to apices of paleae. 


Distribution: §, China (not Taiwan), India, Nepal, Burma, Thailand, N. Vietnam, 
Java, Bali. 


242 Gardens’ Bulletin, Singapore — XXX (1977) 


2. Dryopteris chrysocoma (Christ) C. Chr. Ind. Fil. (1905) 257; Ching, Bull. Fan 
Mem. Inst. Biol. Bot. 8 (1938) 436. —— Aspidium filix-mas var. chrysocoma 
Christ, Bull. Herb. Boiss. 6 (1898) 966. 


Rhizome short-creeping, paleae c. 1.5 cm long, narrow, orange-brown, con- 
colorous, cells very long and slender, margins with minute regular projections. 
Stipe 20 cm X 3 mm, stramineous, bearing broadly ovate paleae 5 X 2.5 mm and 
reduced narrow paleae, Lamina lanceolate, 35 xX 15 cm, bipinnate throughout, 
thick herbaceous, catadromous except at base, both surfaces and axes bearing erect 
capitate orange glandular hairs, rachis beneath also with narrow orange-brown 
paleae. Pinnae linear-lanceolate, basal slightly reduced, 17 pairs, to 9 X 2.7 cm, 
abruptly narrowed to the acute apex. Pinnules c. 12 X 6 mm, sessile but not 
adnate (base narrow), apices broad, rounded, with numerous small teeth. Indusia 
2.5 mm across, persistent, convex, orange-brown, bearing numerous capitate orange 
glandular hairs. 


LUZON. Benguet, Haight’s Place, July 1907, Mearns BS 4219 (MICH). 
Ecology: Terrestrial, c. 2100 m; only one Philippine collection. 
Notes: Very small plants may be fertile. 


Distribution: §. China, N. India & Pakistan, Burma. Reported from Taiwan by 
Tagawa, Acta Phytotax. Geobot. 8 (1939) 22, but not mentioned in the recent 
Flora of Taiwan. 


3. Dryopteris hirtipes (Bl.) O. Ktze., Rev. Gen. Pl. 2 (1891) 813; Christ, Philip. 
J. Sci. 2C (1907) 212; Merrill & Merritt, Philip. J. Sci. SC (1910) 317; Copel. 
Leafl. Philip. Bot. 3 (1910) 806; Fern Fl. Philip. 2 (1960) 281; Holttum, Ferns of 
Malaya, ed. 2 (1966) 635; Sledge, Bull. Brit. Mus. (Nat. Hist.) Bot. 5 (1973) 


6. Aspidium hirtipes Bl., Enum. Pl. Jav. (1828) 148; Christ, Bull. Herb. 
Boiss. 6 (1898) 194. —— Nephrodium hirtipes (Bl.) Hook.; Copel., Polyp. Philip. 
(1905) 21. 


LUZON. Bontoc, Vanoverbergh 3146 (F); Mt Data, Copeland 1887 (MICH), 
Price 389, 395, 402; Mt Pulog, Celestino et al. PNH 3026 (MICH), Curran et al. 
FB 16278 (MICH); Mt Polis, Price 422, 436: Mt Singakalsa, Celestino & Castro 
PNH 3981 (MICH); Mt Santo Tomas, Price 1006, Topping 1187 (NY); Pauai, 
J. K. Santos BS 31667 (US); Benguet, Elmer 6529 (NY, US), Price 1672, Topping 
248 (GH, US), 302 (GH, US), 303 (MICH, US), 3/1 (NY, US); Mt Ragut, 
Loher 850 (US). 


MINDANAO. Davao, Mt Calelan, Elmer 10799 (F, GH, MICH, NY, US); Mt 
Apo, Copeland s.n. 27 Oct 1904 (MICH); Cotobato, Mt Matutum, Copeland s.n. 
1 May 1917 (MICH). 


Ecology: Terrestrial in mountain forest, 1350-2650 m. 


Notes: Dimensions more variable than hitherto reported, in Philippine material 
stipes 12-63 cm, lamina 14-73 cm long, pinnae 2.3-24 x 0.8-3.5 cm. Topping 311 


at NY is aberrant in having irregularly anastomosing veins. Reports of this species. 


from Mt Arayat (Christ 1898, Copeland 1905, 1960) are based on erroneous 


localization of Loher 850, Feb. 1894, by Christ (1898); the specimen in US bears 


the same date and elevation (1950 m) as Christ’s published report but with the 
name Mt Ragut. Arayat is only about 1000 m high. 


Distribution: Sri Lanka and India to S. China & Japan, Malesia, Samoa. 


Philippine Dryopteris 243 


4. Dryopteris wallichiana (Spreng.) Hyl., Bot. Notis. 1953 (1953) 352; Alston, 
Amer, Fern J. 47 (1957) 92; Sledge, Bull. Brit. Mus. (Nat. Hist.) Bot. 5 (1973) 


8. —— Aspidium wallichianum Spreng., Syst. Veget. 4 (1827) 104. —— D. filix- 
mas var. parallelogramma (Kunze) Christ, Philip. J. Sci. 2C (1907) 212; Merrill 
& Merritt, Philip. J. Sci. SC (1910) 317. —— D. paleacea (Sw.) C. Chr. nom. 


illegit.; Copel., Fern Fl. Philip. 2 (1960) 281. 


LUZON. Mt Pulog, Curran et al. FB 16247 (MICH), Jacobs 7260 (A, CAHUP, 
PNH); Mt Data, Copeland 1875 (MICH). 


Ecology: Terrestrial in wet mountain forest, 2100-2800 m. 


Notes: Alston (1957) distinguished the tropical American D. parallelogramma 
from the Asian D. wallichiana by only the color of the rhizome scales, which I find 
to be unreliable. Christensen, Cont, U. S. Nat. Herb. 26 (1931) 280, stated: “‘...a 
close comparison of very complete specimens ... has shown me that it is impossible 
to find one stable character by which they may be distinguished.” The genus 
Dichasium (A. Br.) Fée was established for plants of this species. 


Distribution: Sri Lanka, India, Nepal, Burma, S. China & Taiwan, Japan, Java, 
N. Borneo, New Guinea, Hawaii, Mexico, Cuba, Jamaica, Hispaniola, Guatemala, 
Costa Rica, Panama, Colombia, Venezuela, Ecuador, Peru, Brazil, Bolivia, 
Argentina, Rhodesia, Madagascar. 


5. Dryopteris pulvinulifera (Bedd.)O. Ktze. Rev. Gen. Pl. 2 (1891) 813; Ching, 
Bull. Fan Mem. Inst. Biol. Bot. 8 (1938) 466; Sledge, Bull. Brit. Mus. (Nat. 
Hist.) Bot. 5 (1973) 10. —— Lastrea pulvinulifera Bedd., Ferns Brit. Ind. (1870) 
t. 333. 


Rhizome short-creeping, paleae very narrow, bright golden-brown, glossy, 
c. 1 cm long, entire. Stipe bases swollen, fleshy, blackish, densely paleate; stipes 
to 34 cm long. Lamina deltoid-ovate, acuminate, to 49 cm long, tripinnate + 
pinnatifid at base, anadromous throughout, both surfaces with multiseptate dark 
appressed hairs and unicellular glossy hyaline appressed glandular hairs, rachis 
and costae with lanceolate blackish subclathrate paleae grading distally to bullate 
based. Pinnae to 18 free pairs, basal pinnae to 22 cm long, basal basiscopic 
pinnule to 7 cm long; larger pinnae gradually acuminate, pinnules acute, teeth of 
segments acute (not aristate nor cartilaginous). Sori subterminal on veins, about 
medial, indusia c. 1 mm across, reddish-brown, with multicellular hairs at base of 
sinus and minute fragile appressed glossy glandular hairs elsewhere and at margins. 


LUZON. Mt Data, Copeland 1861 (MICH); Mt Santo Tomas, Price 1616, 1841; 
Mt Burnay, Jwatsuki et al. P-827, P-848, Price 3363. 


Ecology: Terrestrial, rarely petrophytic among mosses, 1800-2200 m, on slopes 
and cliffs in mossy forest. 


Notes: Extremely small fronds may be fully fertile; the smallest, such I have seen, 
is 4.5 cm long including stipe. 


Distribution: Sri Lanka, N. E. India, Nepal. 


6. Dryopteris polita Rosest., Fedde Repert. 13 (1914) 218; Holttum, Ferns of 
Malaya (1955) 492; DeVol & Kuo, Fl. Taiwan 1 (1975) 378. 


Rhizome short-erect, paleae glossy, light brown, subentire, concolorous, to 
13 X 2 mm, Stipe to 52 cm by 3 mm, stramineous. Lamina lanceolate-deltoid, 
18-45 cm long, catadromous above base, bipinnate, texture firm herbaceous, 
beneath with multiseptate appressed reddish-brown hairs, axes with sparse narrow 


244 Gardens’ Bulletin, Singapore — XXX (1977) 


fibrils. Pinnae narrowly lanceolate, acuminate, evenly tapering, broadest on 
basiscopic side; basal pinnae to 17 cm long, stalks 1.6-3.6 cm long, largest basal 
basiscopic pinnule to 6.8 X 1.8 cm, lobed 4/5 to costule basally, most pinnules 
4-9 mm broad with rounded apex. Sori medial, indusia minute, pale brown, com- 
pletely concealed by mature sporangia, sporangial stalks dark reddish-brown 
basally, paraphyses none. 


MINDANAO. Bukidnon, Mt Dumalucpihan, June-July 1920, Ramos & Edano 
BS 39031 (US). 


Ecology: Terrestrial. 


Notes: Rosenstock described this species from Sumatra as exindusiate and an 
isotype in NY is indeed without indusia. However, specimens from Thailand, 
S. Japan, the Philippines, and Papua clearly have very small indusia. In Fern FI. 
Philip. 2 (1960) 283, Copeland cited the one Philippine specimen as an unnamed 
relative of D. sparsa. 


Distribution: Thailand and Sumatra to S. Japan and Papua New Guinea. 


7. Dryopteris sparsa (Don) O. Ktze., Rev. Gen. Pl. 2 (1891) 813; Christ, Philip. 
J. Sci. 2C (1907) 213; Merrill, Philip. J. Sci. 3C (1908) 390; Copel., Leafi. Philip. 
Bot. 2 (1908) 389; Fern Fl. Philip. 2 (1960) 283; Holttum, Ferns of Malaya (1955) 
492, t. 292; Hatusima, Mem. Fac. Agr. Kagoshima Univ. 5 (1966) 22, ——- 
Nephrodium sparsum Don, Prod. Fl. Nepal. (1825) 6; Copel., Polyp. Philip. (1905) 
24. —— Aspidium sparsum (Don) Spreng.; Christ, Bull. Herb. Boiss. 6 (1898) 194. 


BATAN ISLAND. Mt Iraya, Hatusima & Sato 29055 (PNH). 


LUZON. Banawe, Banlugan PNH 72907; Mt Ugo, Ramos BS 5836 (MICH, US); 
Mt Santo Tomas, Price 1042, 1840; Baguio, Price 1578, Topping 222 (GH, US), 
282 (MICH, US); Twin Peaks, Bartsch 211 (US); Benguet, Merrill 7779 (MICH, 
US); Mt Arayat, Loher 1190 (US); Mt Banahaw, Price 362; Mt San Cristobal, 
Price 804, 813, 839; Mt Makiling, Price 211, 366, 499, 2398. 


NEGROS. Mt Cuernos de Negros, Elmer 9949 (F), 10100 (MICH), Herre 110] 
(NY), Price 2612. 


PALAWAN. Foxworthy BS 672 (MICH). 


MINDANAO. Lanao, Clemens s.n. July 1907 (F); Zamboanga, San Ramon, 
Copeland 1727 (MICH, US); Bukidnon, Mt Camates, Ramos & Edario BS 38607 
(US); Davao, Mt Apo, Williams 2526 (NY). 


Ecology: Terrestrial in mountain forests, 800-2100 m, descending to 400 m in 
everwet ravines at base of Mt Banahaw. 


Notes: Occasional specimens have globular sessile yellow glands on the lower 
laminar surface, a character possibly controlled by a single allele. Examples 
are: Herre 1101, Price 813, 1578, 2398. Price 1042 from high elevation on Mt 
Santo Tomas has maroon stipe and rachis. Similar plants have been named D. 
sparsa var, nitidula (Bedd.) Ching, Bull. Fan Mem. Inst. Biol. Bot. 8 (1938) 472. 


Distribution: Sri Lanka & India to S. Japan and Papua New Guinea. 


8. Dryopteris uropinna Price, sp. nov. ——- Aspidium erythrosorum auct, 
non Eaton: Christ, Bull. Herb. Boiss. 6 (1898) 193. —— Nephrodium erythro- 
sorum auct. non (Eaton) Hook.: Copel., Polyp. Philip. (1905) 24. —— D. 
erythrosora auct. non (Eaton) 0. Ktze.: Christ, Philip. J. Sci. 2C (1907) 211; 


Philippine Dryopteris 245 


Copel., Fern Fl. Philip. 2 (1960) 283. —— D. subtriangularis auct. non (Hope 
G Chr. : Ching, Bull. Fan Mem. Inst. Biol. Bot. 8 (1938) 485; Tard. & C. Chr., 
FI. Gén. L’Indo-Chine 7 (1941) 315, fig. 35. —— D. labordei auct. non (Christ) 
C. Chr.: DeVol & Kuo, Fl. Taiw. 1 (1975) 377, p.p. 


Fig. 1 


ce, holotype. A. Frond. B. Stipe scale. C. Pinnule from upper 
Jani Lacy. 


Fig. 1. Dryopteris urop r 
i s ice Glimm 


opinna Pri 
pinna. (see arrow). Drawn by Ms. 


246 Gardens’ Bulletin, Singapore — XXX (1977) 


Species e turma D. erythrosorae, ad D. indusiatam arcte accedens, differt 
magnitudine minore, ambitu subtriangulari, rhachidi glabrescenti, pinnulis late 
rotundati-truncatis, apicibus pinnarum frondisque abrupte contractis et caudatis. 


Ho.otypus: Luzon, Ilocos Norte, Mt Burnay, Price 3364 (MICH). 


Rhizome short-erect, paleae c. 10 X 1 mm, linear, blackish with narrow brown 
margins, entire. Stipes brown, 14-34 cm long, glabrescent above base, Lamina 
deltoid to deltoid-ovate, 17-36 cm long, caudate-acuminate, bipinnate + deeply 
pinnatifid at base, thin papyraceous, dull brown or green, costae stramineous below, 
catadromous above base, above with a few narrow fibrils at axis junctions, below 
with small light brown bullate based acuminate paleae along costae and at costule 
bases, much-reduced septate hair-like paleae along veins. Pinnae to 16 cm long, 
broadest basiscopically, abruptly narrowed and caudate, on stalks to 4 mm long, 
basal pair longest; basal basiscopic pinnule slightly longer or shorter than 
succeeding, 1.5—4.1 cm long, shallowly to deeply lobed. Pinnules blunt, broadest at 
base but not auriculate, when undivided to 20 X 7 mm, with well-spaced small 
sharp teeth. Sori medial or slightly inframedial, indusium orbicular, centrally 
attached, sinus nearly obscured by overlapping edges, glabrous, shriveling but not 
deciduous, to 1.5 mm across, paraphyses short, non-capitate, on sporangial stalks, 
spores dark brown, rather irregularly tuberculate. 


LUZON. Lepanto, Bagnen, Copeland 1929 (MICH); Ilocos Norte, Mt Burnay, 
Iwatsuki et al, P-849, Price 3364. Christ (1898, 1907) also cited a Loher collection 
from Mt Santo Tomas, Feb. 1894, 2250 m. 


CHINA. Kwangsi, Ching 6263 (NY, US); Yunnan, H. T. Tsai 60474 (GH). 
INDIA. Assam, Khasi Hills, 4000 ft., Oct. 1890, G. Mann (NY, US). 
THAILAND. Udawn, Tagawa et al. T-645, T-1297 (US). 

VIETNAM. Cochin-China, Cavalerie s.n. ex Herb. Jeanpert (US). 


TAIWAN. Taito, Tagawa 2670 (MICH); Agyoku, Taihoku, Suzuki 24 (MICH). 
OKINAWA. Mt Yonahadake, Hatusima 18371 (US). 


RYUKYUS. Ishigaki, Koidzumi, s.n. July 1923 (US). 
JAPAN. Kyushu, Yakushima, Iwatsuki 3159 (US). 


Ecology: Terrestrial, 1800-2250 m in sheltered well-lighted spots in moist montane 
forest. 


Notes: Hope cited two specimens of Gustav Mann with his Nephrodium 
subtriangulare, neither of which was seen by Ching (1938) who mistakenly named 
a Clarke specimen as type. Hope’s description differs from Ching’s plant (D. 
uropinna) by rhizome scales castaneous, “‘pinnae all except lowest pair very 
slightly narrowed at base, lowest inferior pinnules on lowest pinnae very much 
shorter than next three or four.” In Philippine literature this species has been known 
as D. erythrosora which however has a creeping rhizome, brown paleae, and much 
reduced basal basiscopic pinnules of basal pinnae, among other differences. D. | 
indusiata differs by attaining larger dimensions, lanceolate outline, more strongly 
paleate stipe and rachis, segments narrower apically, frond apex and pinna apices 
not suddenly contracted and caudate. D. labordei has prominently enlarged basal 
basiscopic pinnules of basal pinnae, brown (not black) paleae, and gradually 
narrowed apices. The specimens above cited from Thailand and Japan differ in 
minor respects from the Philippine type. Some Japanese specimens appear to 


Philippine Dryopteris 247 


intergrade with D. indusiata; they may be hybrids. Another similar species is 
represented by Topping 1794 (US) from Mt Kinabalu; it differs by bullate scales 
absent, segments hardly toothed, basal basiscopic pinnule not dissected. 


Distribution: Himalayan region to S. Japan and Luzon. 


9. Dryopteris varia (L.) O. Ktze., Rev. Gen. Pl. 2 (1891) 814; Ching, Bull. Fan 
Mem. Inst. Biol. Bot. 8 (1938) 490; Copel., Fern Fl. Philip. 2 (1960) 284. 
Polypodium varium L., Sp. Pl. 2 (1753) 1090. —— Aspidium varium (L.) Sw.; 
Christ, Bull. Herb. Boiss. 6 (1898) 192; ibid. II, 6 (1906) 996. Polystichum 
varium (L.) Presl; Copel., Polyp. Philip. (1905) 18. 


LUZON. Bontoc, Madulid et al. PNH 113264, Vanoverbergh 624 (MICH); Ilocos 
Norte, Iwatsuki et al. P-202, P-225, P-886, Price 3342, 3398; Baguio, Copeland 
1959 (MICH), Elmer 6489 (MICH, NY, US), Hernaez CAHUP 13899, 13900, 
13901, Price 386, 1677, Ramos BS 5795 (MICH, US), Topping 177 (US), 
Williams 1513 (NY, US); Mt Banahaw, Price 882. 


Ecology: Terrestrial in seasonally dry forest, 700-1500 m. 


Notes: The Banahaw collection is a new southern range extension. 
Distribution: N.E. India, China, Korea, Japan. 


10. Dryopteris formosana (Christ) C. Chr., Ind. Fil. (1905) 266; DeVol & Kuo, 
Fl. Taiwan 1 (1975) 376. —— Aspidium formosanum Christ, Bull. Herb. Boiss. 
II, 4 (1904) 615. 


Rhizome short-erect, paleae fuscous, concolorous, subentire, to 20 x 2.5 mm. 
Stipes stramineous to brown, to 49 cm long, 6 mm thick at base, paleae blackish, 
entire, Lamina deltoid or deltoid-ovate, to 42 cm long, fully tripinnate at base, 
firm papyraceous, greyish-green, anadromous at proximal base, elsewhere cata- 
dromous. Rachis with reduced narrow blackish paleae, grooves with conical hairs 
at junctions, pinna-rachises, costae, and costules with small dark acuminate bullate- 
based paleae beneath, veins beneath with appressed multiseptate hairs. Basal pinnae 
to 19 cm long, stalk 9 mm, basal basiscopic pinnule notably enlarged, to 13 cm, 
secondary pinnules to 3.5 X 1.1 cm, tertiary pinnules to 8 X 3.5 mm, adnate, 
minutely aristate; basal acroscopic pinnule to 3.2 xX 1.7 cm. Middle pinnae c. 
13 cm long, acuminate, pinnules to 3.4 cm long, secondary pinnules adnate or 
confluent, aristate-denticulate. Sori about medial, indusium bright brown, to 1.3 mm 
across, glabrescent. 


LUZON. Benguet Prov., Mt Santo Tomas. Price 1117; Ilocos Norte Prov., Solsona, 
Price 2929, Mt Burnay, Iwatsuki et al. P-851, Price 3384. 


Ecology: Terrestrial in mountain forest, 1400-2100 m. 
Distribution: Taiwan, Japan. 


11. Dryopteris subarborea (Bak.) C. Chr., Ind. Fil. (1905) 295; Holttum, Ferns 
of Malaya (1955) 491, t. 291; Copel., Fern Fl. Philip. 2 (1960) 284, p.p. 
Polypodium subarboreum Bak., J. Linn. Soc. Bot. 24 (1887) 259. —— D. purpuras- 
cens auct, non (BI.) Christ: Christ, Philip. J. Sci. 2C (1907) 213. 


Rhizome short-creeping, c. 2.5 cm thick, paleae to 20 X 2 mm, brown, con- 
colorous, with marginal projections from one or two adjoining cells bearing a small 
globose pale deciduous gland. Stipe stramineous, sparsely paleate, to at least 55 cm 
Jong, 12 mm thick at base, with c. 12 vascular bundles forming a single ring 
interrupted by the adaxial groove. Lamina to at least 110 cm long, ovate, to fully 
quadripinnate basally, anadromous throughout or nearly so, thin papyraceous, 


248 Gardens’ Bulletin, Singapore — XXX (1977) 


rachis with a few scattered appressed remnants of paleae, rachis groove, pinna- 
rachis grooves, and basal portion of pinnule grooves with minute erect capitate 
glandular hairs within, lamina otherwise glabrous. Basal pinnae to 60 cm long, 
stalk 1.5 cm to basal pinnule, apex acuminate, pinnules to 17 cm long, acuminate, 
secondary pinnules to 4.5 X 1.6 cm, blunt or acute, the proximal pinnate or lobed 
with rounded undulate-toothed segments, grading distally to shallowly crenate- 
dentate. Middle pinnae c. 32 cm long, acuminate, stalk 1 cm, basal pinnules 
7.5 X 3 cm, secondary pinnules to 1.6 X 0.9 cm. Sori medial to slightly supramedial, 
indusium brown, glabrous, smaller than sorus, 0.5 mm across, paraphyses with 
hyaline or yellow bead-shaped heads. 


LUZON. Quezon, Real. Llavac, Price 1494, 1499; Laguna, Mt San Cristobal, 
Copeland PPE 91 (GH, MICH); Mt Makiling, Elmer 18067 (F, GH, NY), 
Price 2988. 


MINDORO. Mt Halcon, Merrill 6101 (MICH)? 
CATANDUANES. Ramos BS 30192 (US), 30223 (US), 30462 (US). 
SAMAR, Gandara, Price & Hernaez 158. 


PANAY. Capiz, Mt Bulilao, Martelino & Edafio BS 35645 (US), 35652 (US), 
35693 (US), 35730 (US). 


Ecology: Epiphytic on trunks and large branches in wet forest, 300-1200 m. 


Notes: I place here with doubt Merrill 6101 known to me by a single sterile 
specimen pustular beneath. According to the field label, a Bornean specimen at 
Kew, matching the type, was also epiphytic with short-creeping rhizome (R. E. 
Holttum, in litt.). Related is the New Guinean D. pseudoparasitica v.A.v.R. with 
relatively slender and long-creeping rhizome and dimorphic fronds. 


Distribution: Sumatra, Malaya, Borneo (Java?). 


12. Dryopteris purpurascens (B].) Christ, Philip. J. Sci. 2C (1907) 213, nomen 
tantum; Copel., Leafi. Philip. Bot. 3 (1910) 807; Backer & Posthumus, Varenflora 
voor Java (1939) 47, p.p. Aspidium purpurascens Bl., Enum. Pl. Jav. (1828) 
169. —— Nephrodium divisum auct. non Hook.: Copel., Polyp. Philip. (1905) 
26. —— D. subarborea auct. non (Bak.) C. Chr.: Christ, Philip. J. Sci. 2C (1907) 
214, p.p.; Copel., Leafl. Philip. Bot. 3 (1910) 807; Fern Fl. Philip. 2 (1960) 
284, p.p. 


MINDANAO. Mt Apo, Copeland 1138 (MICH, NY, US), 16/4a (MICH), s.n. 
5 Sept. 1932 (MICH), Elmer 11418 (MICH), 11550 (F, GH, MICH, NY, US), 
11842 (F, GH, MICH, NY, US), Williams 2479 (NY, US); Mt Matutum, Cope- 
land s.n. 30 April 1917 (MICH), Ramos & Edafnio BS 85302 (GH). 


Ecology: Terrestrial, petrophytic, or epiphytic on mossy trunks, 1500-2000 m. 


Notes: Elmer 11842 has a different appearance because of maroon color when 
dry and finer teeth of segments; it may represent another species. Mindanao 
specimens which I am treating as D. purpurascens differ from the next species, D. 
permagna, by being somewhat laxer and less dissected with broader usually deltoid 
teeth and inconsistently catadromous or anadromous architecture, as well as the. 
paraphyses. The holotype of Aspidium purpurascens Bl. (photo MICH, US, Morton 
neg. 1304) consists of two separate pinnae evidently from the same frond. labeled 
as collected in the Moluccas by Zippelius. Blume’s variety Beta (photo MICH, US, 
Morton neg. 1305) seems to be identical, possibly parts of the very same frond, 
consisting of a frond apex and the basal portion of a basal pinna, but is labeled 
Java, collector Blume. Both are quadripinnate, and anadromous throughout. While 


Philippine Dryopteris 249 


no Philippine materials are an exact match to the type of D. purpurascens, I refrain 
from describing the Mindanao plant as new without further information. Among 
other subtle differences, Javan plants have relatively numerous persistent narrow 
paleae along axes below. 


Distribution: Java, Celebes, Moluccas (Borneo?). 


13. Dryopteris permagna Price, sp. nov. —— D. subarborea auct. non (Bak.) 
C. Chr.: Christ, Philip. J. Sci. 2C (1907) 214, p.p.; Copel., Fern Fl. Philip. 2 
(1960) 284, p.p. 

Rhizoma crassum, erectum, paleis usque ad 42 X 7 mm cellulis marginalibus 
aliquot eminentibus ferens. Stipites usque ad 103 X 1.2 cm. Lamina subquinque- 
pinnata, ovata, usque ad 138 cm longa, herbacea. Pinnulae primariae pleraque in 
positione catadromica portatae. Pinnae basales amplissimae, usque ad 64 cm 
longae, petiolus usque ad 10 cm longis, pinnula basali basiscopici usque ad 
31.5 cm longa. Pinnulae ultimae apice rotundatae denticulatae, lateribus lobatae vel 
dentatae, lobis acutis erectopatentibus usque fere patentibus. Costae, costulae 
venaeque infra pilis persistentibus multiseptatis appressis praeditae. Sori medialis, 
indusia 0.5-1 mm diametro, glabra. Paraphyses inconspicuae, hyalinae, non 
capitatae, 

HoLotypus: Luzon, Mt Santo Tomas, Price 16J]] (PNH). 


Rhizome stout, erect. Paleae to 42 X 7 mm, long-acuminate above a relatively 
broad lower portion, brown, concolorous, margins with small blunt projections 
consisting of the outcurved portion of one cell or two adjacent cells. Stipe light 
brown, with smaller narrower paleae, to 103 cm long and 12 mm thick, with 15 
vascular bundles near base, the two adaxial on either side of the stipe-groove much 
the largest. Lamina ovate, to 138 cm long, olive-green, herbaceous, quadripinnate or 
subquinquepinnate at base, pinnae and major pinnules long acuminate, most or all 
pinnae with basal primary pinnules in catadromic position, secondary pinnules in 
anadromic position except sometimes for the larger ones of basal pinnae, tertiary 
pinnules and quaternary segments nearly all in anadromic position. Major axes 
beneath with deciduous narrow fibrils, persistent appressed reddish-brown multi- 
septate hairs c. 0.3 mm long borne along costae, costules and veins beneath, 
sometimes a few on upper surface. Basal pinnae largest, to 64 cm long on stalks 
to 10 cm long, basal basiscopic pinnule to 31.5 cm, stalk to 1.8 cm, not very 
much larger than the succeeding, basal acroscopic pinnule to 20.5 cm, very slightly 
shorter than the several succeeding. Largest secondary pinnule 10 cm long, tertiary 
pinnules to 2.5 X 1.7 cm, largest quaternary pinnules broadly adnate or connate, 
to 10 X 5 mm. Third pair of pinnae from base 45 cm long, fifth pair 33 cm long 
with stalk 2 cm, tenth pair 12 cm long, ultimate pinnules with broad blunt 
denticulate apices and spreading-ascending pointed lobes or teeth. Sori about medial, 
indusia 0.5-1 mm across, brown, glabrous, paraphyses shorter than sporangia, 
apices hyaline, not or hardly expanded. 


LUZON. Mt Polis. Oliver PNH 30576; Mt Data. Ramos & Edario BS 40285 (US): 
Mt Pauai, Sulit PNH 5120 (MICH): Bontoc, Ramos & Edanio BS 37834 (US), 
37847 (US), 37985 (US), 38064 (US); Mt Ugo. Ramos BS 5777 (MICH, NY, 
US); Haight’s Place, Mearns BS 4194 (MICH, US); Baguio, Loher 832 (US); Mt 
Santo Tomas, Price 1611, 1862. 


Ecology: Terrestrial in mountain forest, 1400-2200 m. 


Notes: Ramos & Edafio BS 37847, very congested and nearly glabrous, is placed 
here with some doubt. This species, apparently endemic to Luzon, is very similar 
to Polynesian specimens of D. arborescens (Bak.) O. Ktze., which is characterized 
by broad pale brown paleae on costae, the paleae attached parallel to the costa. 


250 Gardens’ Bulletin, Singapore — XXX (1977) 


Another similar Polynesian plant, D. maxima (Bak.) C. Chr., is anadromous, 
very finely dissected, with supramedial sori. Some New Guinean specimens 
are similar to those of Polynesia; none are quite the same as anything in the 
Philippines. 


Acknowledgements 


I am grateful to the directors and curators of the following institutions for 
the use of facilities, loan of specimens, and other forms of assistance: A, C 4 
F, GH, KYO, MICH, NY, PNH, SING, TAI, US. I am especially indebted to 
Prof. T. J. Rimando of the University of the Philippines at Los Bafios for much 


help when collecting in the Benguet highlands of Luzon. 


Fertile-sterile Leaf Dimorphy in Ferns 
Warren Herb Wagner, Jr. and Florence S. Wagner! 


Department of Botany, University of Michigan 
Ann Arbor, Michigan, U.S.A. 


In addition to his many other contributions, Dr. R. E. Holttum has been 
unquestionably one of the world’s leading authorities on the taxonomy of ferns with 
fertile-sterile leaf dimorphy, especially the lomariopsid group. Inspired in part by 
reading his papers, we became interested several years ago in the over-all subject 
of fertile-sterile dimorphy and began the study to be reported here. As will be 
seen, there are numerous disagreements in the interpretations of dimorphy, and 
this represents an effort to review these as well as other aspects of the general 
phenomenon. 


Dimorphy of fertile and sterile fronds is widespread and bears upon both the 
systematics and ecology of ferns. It is an expression of a trend found throughout 
vascular plants, a trend which achieves its ultimate in seed plants. Recognizing 
that the term “‘dimorphy” may sometimes apply to other than fertile-sterile (i.e., 
various types of heterophylly of vegetative leaves in ferns), for convenience the 
term here will be confined to differentiation into sporophylls (spore-producing 
fronds, fertile Blatter) and trophophylls (purely photosynthetic fronds, Laub- 
blatter). Profound alterations may appear in the evolutionary transition from 
primitive monomorphy, including transformations in morphology, anatomy, physio- 
logy, and ecology. These changes are thought to possess adaptive value. However, 
this idea is not universally accepted, and there is little or no experimental evidence 
to support it. At present, in fact, most of our conclusions are intuitive and hypo- 
thetical, and we strongly need rigorous tests. We hope that the ideas we present 
here will help to stimulate such experimentation. 

As many as one in five of all fern species possess what may be called fertile- 
sterile dimorphy, but this figure has to be arbitrary because of so many transitions 
that occur between monomorphy and dimorphy. Where the dimorphic condition 
is strongly expressed, taxonomists have traditionally utilized it as a useful character 
in species discrimination. In some instances, taxonomic separations have been 
created even at the generic level on the basis of sporophyll-trophophyll dimorphy. 


The analysis of foliar dimorphy invokes a multitude of biological questions 
such as what parts undergo changes, what are the effects on sporangial arrangement, 
what developmental processes underlie formation and timing of fertile expressions, 
what — if any — ecological advantages are conferred, and to what extent 
dimorphism is valid as a systematic character, The following report is an outcome 
of our observations in the field, laboratory, and herbarium, together with a survey 
of the literature. The most thorough and most recent survey seems to be that of 
F. G. Dickason in 1946. Because of the widely scattered nature of references to 
this subject, we fully expect that some important items have been overlooked. We 
also anticipate that various ideas expressed here will be subjected to future 


1 Research carried out in connection with National Science Foundation Grant DEB-75- 
03550, ‘Evolution and Taxonomic Applications of Foliar Characters.” Various individuals 
and institutions made available specimens for this study, especially Michael G. Price. Janice 
Glimn Lacy provided many of the microscope preparations and the diagrams, and various 
individuals aided in the development of this manuscript. To all we express our warm thanks. 


251 


252 Gardens’ Bulletin, Singapore — XXX (1977) 


reappraisal, If, however, this article serves to stimulate critical research, its purpose 
would be fulfilled. 


All degrees of morphological and anatomical divergence exist between fertile 
and sterile fronds. In fact, one of the reasons the term “‘monomorphic”’ is rarely 
used may be that practically all ferns are strictly speaking at least slightly 
dimorphic. Steps in dimorphic divergence are well exemplified in numerous genera. 
In eastern North America, for example, we encounter a good illustration in the 
woodfern genus, Dryopteris. D. goldiana (Hook.) Gray has hardly any detectable 
difference between soriferous and non-soriferous mature fronds. D. cristata (L.) 
Gray, with erect, tall fertile fronds, and spreading, short sterile fronds, shows 
strong differentiation. D. spinulosa (O. F. Muell.) Watt is intermediate, Other good 
examples of such sequences are met with over a broad taxonomic spectrum, such 
as, in Polypodioideae, Loxogramme, Microgramma, and Pyrrosia; in Blechnoideae, 
Blechnum; and in Dryopteridoideae, Davallia, Humata, Nephrolepis, Oleandra, 
and Polystichum?. 


It should be understood that the term ‘“‘dimorphism”’ does not necessarily 
imply a precise morphological parallelism. The changes involved are probably 
often the result of analogous, rather than homologous, transformations, and different 
examples show different degrees of involvement of petioles, blades, pinnae, 
sporangial arrangements, histology, and physiological reactions. 


The sporophyll-trophophyll differentiation may embrace whole fronds or only 
parts of fronds, and in the latter, it may be the tip that is fertile, the middle of the 
frond, or the base. These modifications are diagrammed in Figure 1. Interfoliar 
dimorphy, constituting complete differentiation of whole fronds is the most 
common type. However, because of the commonly transitional nature of the 
manifestations, many fertile fronds are merely “‘subdimorphic” (term as used by 
Dickason, 1946) in the sense that the changes displayed are incomplete in kinds 
and amounts. A “holodimorphic”’ situation includes total loss of vegetative function 
in the sporophyll. However, the latter is hard to achieve because during the early 
stages of maturation the rachises and costae of all fertile fronds (at least those 
known to us) are to some extent green and photosynthetic. Whole-frond differentia- 
tion into sporophylls may follow two routes. It may be the culmination of an 
evolutionary trend in which at first only the frond tip is fertile. Dickason (1946) 
suggests that there is a progressively earlier shift from the vegetative to the 
reproduction condition of the leaf starting at the apex (i.e., Fig. 1, B to D) mediated 
by an hypothetical “‘sporogen” growth substance. On the contrary, one can just 
as well imagine modification of the whole leaf progressing from the inception of 
the leaf, bottom to top. 


Where fertile-sterile differentiation appears in separate parts of the same leaf, 
“the term hemidimorphism is used instead of dimorphism’’ (Dickason, 1946). The 
hemidimorphic leaves with terminal fertile pinnae are the most widespread. In 
Osmundaceae, this condition is illustrated by the familiar O. regalis L.; in Poly- 
podioideae, by the genera Paltonium and especially Belvisia; in Grammitidoideae, 
by Grammitis Sect. Xiphopteris; and in Adiantoideae, by Llavea. Two extremes in 
the Dryopteridoideae are observed in the eastern North American Polystichum 
acrostichoides (Michx.) Schott and Dryopteris ludoviciana (Kze.) Small. As in 
other forms of hemidimorphism, the fertile pinnae even when they are 
photosynthetic at the time of spore production as they are in D. ludoviciana ——. 


are shorter-lived than the sterile, and turn brown and die well before the sterile 
pinnae. 


Examples of hemidimorphism in which only the medial pinnae are the fertile 
ones are known best in Osmunda, and most familiarly in O. claytoniana L.. which 
is designated in North America by the colloquial name of “‘interrupted fern” 


2 The family-subfamily classification used in this paper is that of W. Wagner (1973). 


Leaf Dimorphy in Ferns 253 


because its fertile pinnae discharge their spores and fall off early in the season 
(usually by early or middle July), leaving fronds in which the lower and upper 
regions of photosynthetic pinnae are set apart by a length of naked rachis. 


Ferns with the hemidimorphic condition effecting only basal pinnae are 
taxonomically a motley lot. In the striking cyatheoid genus Thyrsopteris, known 
in the living flora only from Juan Fernandez Islands, several pairs of lower pinnae 
are fertile, these extremely skeletonized (i.e., lacking lamina) but bearing sori with 
conspicuous cup-like indusia. More abundant ferns with modified basal pinnae 
occur in Ophioglossaceae and Schizaeaceae subg. Anemioideae. In the genus 
Anemia we find a spectrum running from only subtly subdimorphic basal pinnae 
all the way to fully differentiated basal pinnae. Accompanying this progressive 
laminar reduction and skeletonization of the basal pinnae pair, there is increased 
tendency for them to become erect and tower above the spreading sterile pinnae. 
In the most evolved forms in Anemia, the sporangial masses are held well above 
the photosynthetic portions of the leaves, and effectively the same result is achieved 
as in those dimorphic leaves in which the sterile fronds are flat and form rosettes 
(cf. Fig. 1, C, hemidimorphic, with Fig. 2, D, holodimorphic). The alterations in 
Anemia have been described in some detail by Mickel (1962, 1967). One of the 
interesting features in specialized members of this genus is the close association 
of the fertile basal pinnae in point of apparent origin with the second pair of 
pinnae, i.e., the lowest of the sterile pinnae, a condition to which Mickel refers as 
“fertile pinnae approximate to sterile.” On the basis of comparative morphology, 
we can conclude that the fertile petiolule bases have fused with the main petiole 
up to the first sterile petiolules. A parallel condition is seen in the single fertile 
spike of certain Ophioglossaceae, e.g. Botrychium virginianum (L.) Sw. 

Mickel (1967) describes the probable evolution of the fully fertile fronds of 
certain species of Anemia as following a pathway opposite that suggested by 
Dickason (see above) for other holodimorphic ferns. Most Anemia taxa produce 
a single pair of fertile basal pinnae, but in A. rutifolia Mart. there is more than 
one pair of basal fertile pinnae, and in A. millefolia Gardn. all of the pinnae are 
fertile. We do not believe that the two models are necessarily in conflict, nor the 
idea, suggested earlier, that in some cases the sporophyll may have been holo- 
dimorphic from the beginning. There are thus three different possible routes: (a) 
change directly from monomorphic leaves to holodimorphic (Fig. 1, A-D); (5) 
change from monomorphic to hemidimorphic, with the fertile pinnae apical, to 
holodimorphic (Fig. 1, A-B.D); and (c) from monomorphic to hemidimorphic 
with the fertile pinnae basal to holodimorphic (Fig. 1, A-C-D). Monographers 
should be prepared to encounter any one of these routes in a given evolutionary 
line. 


Morphologically the single fertile spike of Ophioglossaceae (Botrychium, 
Helminthostachys, Ophioglossum) resembles the fertile basal pinna condition in 
Anemia, differing from it in the apparent fusion to form a single primordium and 
mature structure. This interpretation ——— the ‘“‘Fusion Theory” or “‘Peltation 
Theory’’ —— to explain the fertile spike, is the most parsimonious and requires 
the least number of assumptions *. Unfortunately, the morphological intermediates 
that would directly confirm the phylogeny of the fertile spike have been lost from 
the fossil record. All we have are data from living forms to support this interpreta- 
tion ——— the pattern and origin of vascular strands, and the occasional grapefern 
specimens (especially in Botrychium subg. Sceptridium, considered primitive on 
other grounds) of the Anemia condition with separate fertile basal pinna pairs, as 


3In the welter of interpretations to which the fertile segment of Ophioglossaceae has been 

subjected, one of the most unusual is that of Wieffering (1964), who, regarding Ophioglossum, 

treats the sterile segment of the frond as a “trophophyll’ and the fertile spike as comprising 
a “fertile stalk” terminated by a “strobilus”! 


254 Gardens’ Bulletin, Singapore — XXX (1977) 


well as intermediates, There are a few examples of adder’s-tongue ferns, Ophio- 
glossum, with the sterile blade reduced or suppressed entirely. However, the belief 
that the rare South African species, O. bergianum Schlecht. has complete sporophyll- 
trophophyll separation (as believed, for example, by Copeland, who keyed it out 
on this basis in 1947) has proved to be erroneous. In our own observations of this 
curious plant we find an illusion of two separate fronds that is created by a very 
low attachment of the fertile spike to the sterile segment. 


Although the details are not agreed upon, the consensus seems to be that the 
sporocarps of the Marsileaceae are modified basal pinnae in which the sporangia 
are encapsulated. Unless it can be established that the sporocarps are de novo in 
origin, comparative morphology dictates that the homology to basal pinnae is the 
most likely explanation of these very distinctive structures, especially if it is agreed 
that Marsileaceae are specialized leptosporangiate ferns. Gupta (1956, 1962) has 
brought together much of the information bearing upon the nature of the sporocarp. 
Also, he has found (pers. comm.) very interesting forms of Marsilea minuta L. 
which have numerous sterile pinnae borne along the leaf axis, suggesting a reversal 
to a more primitive condition. It is easy to speculate that the hemidimorphic 
condition of Marsileaceae may have more than a coincidental resemblance to that 
of the Anemioideae. 


Attention should be called to the less spectacular but substantial changes that 
often occur in the specialization of distinct trophophylls. The obviously striking 
aspect of many sporophylls quickly attracts attention from morphologists and 
taxonomists, but often the trophophylls themselves do not remain merely as sterile 
remnants but little changed from the original double-duty leaves of the mono- 
morphic ancestors. The uncommon curly-grass fern, Schizaea pusilla Pursh, of 
northeastern North America has trophophylls which are wiry, twisted organs 
concentrated at the base of the plant. In various polypodioid genera, such as 
Microgramma in the New World and Pyrrosia in the Old, there seems to be an 
inverse correlation between changes in the sporophyll and those in the trophophyll: 
As the sporophyll becomes longer and narrower, the trophophyll becomes shorter 
and wider. Holttum (1954) illustrates Pyrrosia nummularifolia (Sw.) Ching, an 
extreme example in its genus with the sterile fronds “‘very shortly stalked, almost 
circular to broadly ovate.” In this example, the fertile fronds are more similar to 
the norm of the non-dimorphic or subdimorphic fronds of the genus than are the 
sterile. Figure 2, D, illustrates this type of change in trophophylls. 


In a few of the paddle-ferns, Elaphoglossum, the trend described above is 
reversed: The trophophylls become linear, extremely long and narrow, and pendent, 
while the sporophylls retain their lanceolate outlines. Splendid illustrations of such 
specialized hanging trophophylls are the New World tropical Elaphoglossum 
siliquoides (Jenm.) C. Chr. and especially E. herminieri Bory ex Fée. 


In teleological terms, the “‘need’’ for certain leaf shapes required by carrying 
out both duties of photosynthesis and sporulation seems to be eliminated if 
sporophylls evolve, permitting the trophophyll to become more adapted to its role. 
In Dryopteris cristata, the sterile leaves are smaller and more spreading, so that they 
form a “‘light-catching” rosette at the base of the plant in contrast to the bulk of 
the species of Dryopteris in which the sterile fronds are less differentiated from the 
soriferous fronds. In general, in terrestrial forms, the flatter and rounder the 
trophophyll, the more efficient it will be photosynthetically. This seems to be true 
also in those epiphytic ferns, such as the creeping polypodioids, in which the 
trophophylls “hug” the branches and twigs of the host trees and shrubs. 


A somewhat neglected component of comparative morphology of the sporophyll 
is the modification of sporangial distribution. The over-all trend is toward the 
concentration of sporangia, but this concentration is achieved in a variety of ways. 
The sori may keep their organization and simply become more tightly grouped. 


Leaf Dimorphy in Ferns 255 


oe 
a( Hh 


Figure 1. Diagrammatic representation of fertile-sterile dimorphy. 4. Prmitive condition: 
monomorphic. D, F. Complete dimorphism of sporophyll and trophophyil, showing different 
relative sizes of sporophylls. B,C, E. Hemidimorphic conditions: B Fertile parts terminal 
E. Fertile parts medial. C. Fertile parts basal. 


They may, on the other hand, lose their soral organization and fuse into a con- 
tinuous “‘sheet” of sporangia spread over the sporophyll lamina, or the entire 
sporangial area may become skeletonized, losing lamimar webbing completely and 
producing clusters of single-veined axes. The sporangia may be exposed, even m 
the youngest stages of development (as in certain acrostichoid ferns). or they 
may be protected by indusia, paraphyses, or by inrolling of laminar tissue to form 
more or less persistent sheltering capsules. Also, these various modifications may 
be combined in different ways. 


Compacted but still discrete sori are observed in diverse taxonomic groups. 
The exposed epidermal areas between the sori are reduced so that finally no bare 
can be seen and the sori are in contact on all sides. For polypodioid sori 

this is well illustrated in the genus Pyrrosia. While it is true that after spore 
discharge members of this genus may appear to have truly acrostichoid soriation, 


256 Gardens’ Bulletin, Singapore — XXX (1977) 


A 


Figure 2. Diagrammatic representation of relative differentiation of sporophylls and 
trophophylls, A. Primitive condition: monomorphic. B,C, D, Dimorphic. C. Extreme change 
in sporophyll, the characters (height, orientation, exaggerated). D. Extreme change in 
trophophyll, specializations of size, shape, and orientation strong. 


this is actually an illusion. Sori of the ancestors of Pyrrosia were probably of the — 
type seen today in Pleopeltis well separated, exindusiate, and round. An 
outstanding example of nearly confluent indusiate sori is present in the eastern 
North American Polystichum acrostichoides (L.) Schott. Here the situation 
resembles that in Pyrrosia in that after spore discharge the fertile pinnae become 
falsely acrostichoid, Prior to spore discharge, however, not only are the discrete 
sori recognizable, but each is provided with a conspicuous peltate indusium. 


Leaf Dimorphy in Ferns 257 


Truly acrostichoid soriation results from the origin of sporangia upon a 
continuous placental surface which extends across the abaxial surface of the blade 
or pinna. Even in young stages it is not possible to make out individual sori. 
Although there are some disagreements over details of interpretation and termi- 
nology (cf. W. Wagner, 1965, pp. 299, 300). recognition of this type of soriation 
is generally accepted as a valuable tool in generic taxonomy. Ferns with acrosti- 
choid soriation may or may not have laminar contraction associated with it. In 
Elaphoglossum (Aspleniaceae subg. Dryopteridoideae) and Acrostichum (Adianta- 
ceae), the fertile blades or pinnae may be only slightly contracted or not contracted 
at all. However, they tend to be more ephemeral than the sterile blades or pinnae, 
becoming curled and dry before the sterile segments. Acrostichoid soriation in 
unrelated taxa probably has had different modes of origin. In Acrastichum the 
ancestors were very likely gymnogrammeoid ferns lacking indusia. The sori were 
linear and followed the veins, and they were separated by exposed laminar surface. 
By extension of the sori into the naked areas and finally becoming confluent, the 
acrostichoid condition came about, the sporangia making up a continuous sheet, 
clothing the entire laminar undersurface. On the contrary, the history of Elapho- 
glossum may have followed a very different route. If we accept the comparative 
evidence from many characters, Elaphoglossum is a derivative of ferns with round, 
indusiate sori (like those of Dryopteris and Polystichum). The steps in the evolution 
of Elaphoglossum from its probable progenitors may then be suggested to have 
followed this route: (1) assumption of the epiphytic habitat, (2) simplification 
of the frond, divided to undivided, (3) approximation of sori, from remote to 
continuous (as seen today in Polystichum acrostichoides). (4) loss of indusia, and 
(5) unification of placental tissue all across the laminar surface to become acrosti- 
choid. Some of these steps may have taken place in a different order. The 
important point is that the acrostichoid condition may be the result of convergent 
evolution and not necessarily be a signal of systematic affinity. Surely the examples 
known in the polypodioid ferns (e.g., Belvisia, Christiopteris, Dendroglossa, and 
Platycerium) arose from very different ancestors from those given above. 


Bower pointed out long ago (1923) that some acrostichoid ferns have “no 
special vascular provision for the nutrition of the sporangia”, whereas others 
develop elaborate modifications involving two parallel vascular systems ——— the 
normal venation plus a specialized receptacular venation on the abaxial side. The 
two are in connection with each other, of course. The state of having two parallel 
vascular systems is termed “diplodesmic”, and is especially well shown by 
acrostichoid examples in the Polypodioideae (Bower, 1928). 


What we here call the botryoidal condition, in which the sporangia form 
clusters upon more or less naked and branched axes, is best shown in three groups 
—Ophioglossaceae, Schizaeaceae subg. Anemioideae, and Osmundaceae. Here 
the flat laminar condition is partially or entirely lost, and the sporangial masses 
become three-dimensional and glomerulate. Assuming that lamina-borne sporangia 
are ancestral (see discussion below) we may conclude that the primitive condition 
in Osmundaceae is that displayed by Todea and in Anemioideae by Mohria or 
certain members of Anemia itself. The only presumed primitive or intermediate 
conditions found in Ophioglossaceae are occasional forms or terata in which the 
sporangia are borne upon partly or fully developed lamina. This is especially well 
shown in certain species (e.g., Botrychium matricariifolium A. Br. and B. simplex 
E. Hitche.) in which a percentage of every population shows sporangia arising 
from the sterile segment margins. 


Sporophylls which have the encapsulated or angiosporangial condition are 
especially well developed in Plagiogyriaceae and Aspleniaceae subf. Blechnoideae. 
In these ferns the sporangia are covered, at least partially, by the inrolling of the 
pinna margins. They are protected until they are fully mature and ready to 
discharge the spores. To study early stages of sporangial development it is necessary 


258 Gardens’ Bulletin, Singapore — XXX (1977) 


to cut away or pull back the folded-over lamina. The most highly evolved 
encapsulated structures are those of the onocleoid ferns, especially Matteuccia and 
Onoclea, in which the sporophyll capsules do not open until months after the 
sporangia have matured. 


Fertile-sterile differentiation at the histological level has been largely uninvesti- 
gated and we are currently trying to determine the scope of the anatomical changes 
associated with sporophylls. By using the frequent intermediates that occur as 
abnormalities in many dimorphic taxa, it is possible to detect gradients and 
establish homologies. Techniques of clearing and staining portions of the leaves 
are especially suited to this type of study. A brief summary of our observations 
is as follows: Narrowing of the segments of sporophylls is accomplished by the 
progressive reduction of the lamina until, in extreme cases like the botryoidal 
types described above, the axes become terete and there is no trace of laminar wings. 
Associated with this there is a large increase in the ratio of over-all surface to 
volume, allowing for more rapid drying. With reduction in lamina we usually 
notice an increase in the width of the vein intervals and a simplification of the vein 
pattern. The latter is accomplished by eliminating some of the vein branchings 
and most or all of the areoles, as in Lorinseria areolata (L.) Pres] (Blechnoideae) 
and Onoclea sensibilis L. and Onocleopsis hintonii F, Ballard (Dryopteridoideae, 
Labouriau, 1958; Lloyd, 1971). In Lomariopsis the sporophylls have free-ending 
veins, lacking the marginal connections of the trophophylls (Holttum, 1939). 
Comparative investigations of the vein intervals of nearly two dozen taxa were 
made by Lommasson and Young (1971), but they did not compare conditions in 
sterile and fertile fronds. 


There may be simplification in other laminar tissues as well. The epidermis 
may be softer in sporophylls than in trophophylls, and may tend to produce fewer 
stomates. The epidermal cells may be larger and straighter-walled. The mesophyll 
may have less elaborate spongy parenchyma that is composed of larger, thinner- 
walled cells. In most strongly dimorphic taxa, the cells lose their chlorophyll at 
maturity, appearing en masse a whitish, yellowish, or brownish color. Where the 
acrostichoid condition occurs, the sporangial initials are no longer localized in 
punctate or linear receptacles, but become spread over the abaxial surface, forming 
a continuous placental area both along and between the veins. As indicated earlier, 
the sporophyll venation of certain acrostichoid ferns may actually be more complex 
than that in the trophophylls. 


The physiological study of sporophyll-trophophyll differentiation is still in its 
infancy. One thinks of such names as Labouriau, Sharma, Steeves, Sussex, Wardlaw, 
and Wetmore in connection with processes of initiation and development of fertile 
leaves. White (1971) has reviewed much of their work. There seems to be little 
question that formation of sporophylls is controlled hormonally, and that their 
timing is related to periods of light and perhaps heat, although there is a vast 
amount of research needed to clarify the factors involved. Various extraneous 
influences may interfere with the normal course of sporophyll development, causing 
intermediate fronds to form. There are scattered reports on effects of injuries 
brought on by such agents as fire, flood, mowing, destructive substances (e.g., tar), 
and the trauma of transplanting, in inducing intermediate fronds, but there are 
very few controlled experiments to warrant general conclusions, It does seem that 
destruction of many or all of the actively photosynthetic fronds of a given plant 
will have a tendency to produce imperfect expressions of sporophylls in the new 
crop of fronds. In Polystichum acrostichoides, the partially fertile, subdimorphic 
pinnae of “forma incisum (Gray) Gilbert’? apparently result from abnormal new 


growth of late leaves after injury of the normal fronds of the season (Wagner, 
Farrar, and McAlpin, 1970). 


Leaf Dimorphy in Ferns 259 


In very strongly marked dimorphy, the fertile frond becomes less responsive 
to light and more responsive to gravity. The typical spreading orientation of the 
trophophyll in which the blade approaches a position perpendicular to the direction 
of light is absent in the sporophyll which is now so negatively geotropic that it 
becomes totally erect. A simple and quick experiment to demonstrate tropism of 
fertile parts may be made with Botrychium virginianum (L.) Sw. If whole leaves, 
including both the fertile and sterile segments, are cut off at ground level during 
the active growing season before the sporangia have fully matured or dicharged, 
and laid in pans of water in various positions, we have found that at room tempera- 
ture in only 10 to 20 hours all of the fertile spikes will twist and become upright, 
no matter what the position of their associated sterile segments, illustrating dramati- 
cally the “‘urge”’ to raise the sporangia to the highest position possible. 


The tissues of the sporophylls, being simpler than those of trophophylls and 
having a more flimsy structure, are more readily dried out. Their vertical position 
in relation to sun and wind, in contrast to the trophophylls which are closer to the 
substrate and more protected, also promotes dessication, Presumably the sporophylls 
have “‘cheaper construction” partly because they are more ephemeral and, lacking 
photosynthesis, have no value to the plant after their function of spore production 
is finished. Support by the rest of the plant is withdrawn when the sporophyll’s job 
is done, and the fertile frond accordingly turns brown, dries out, and falls away. 
This is shown especially well in single hemidimorphic leaves. The differentiated 
fertile tips of Osmunda regalis and Polystichum acrostichoides dry up and die long 
before the sterile parts of the same leaf. In Botrychium, Ophioglossum, and Anemia, 
the photosynthetic pinnae continue to function as if nothing had happened, even 
though major parts of the leaf, the fertile segments, have collapsed and turned to 
twisted debris. 


The seasonal timing of sporophylls and trophophylls may differ profoundly 
in taxa occurring in the same geographical areas and habitats. Nozu (1968) studied 
periodicity in Japan of members of nine genera with respect to sterile-fertile frond 
maturation, and reported that in a given species they may differ by as much as six 
months, Usually the sterile fronds mature first during the course of the year, but 
in Osmunda japonica Thunb. the fertile fronds mature a month or so earlier than 
the sterile. 


A most curious state of affairs is found in the periodicity of Botrychium 
lunarioides (Michx.) Sw., a rarity of the southeastern United States known as 
‘Winter Grapefern.” The frond arises above ground in November and the sterile 
segment expands to its full size by sometime in January. The fertile segment, on 
the contrary, remains curled over at the ground level and stays soft and meriste- 
matic for nearly three months, only expanding and become erect in March. Spore 
discharge occurs in late March and then the entire leaf, including both fertile and 
sterile segments, dies in April. The plants are dormant, and completely buried 
under ground, from April to November. 


Of what adaptive value is fertile-sterile leaf dimorphism? It is found in plants 
of such widely different habits and habitats that it is practically ubiquitous. The 
condition is found in giant ferns (e.g., Acrostichum) and midget ferns (e.g., 
Peltapteris), in thick-textured ferns (e.g., Polystichum) and thin-textured ones (e.g.. 
Trichomanes). Dimorphy occurs in ferns with simple leaves (Pyrrosia) and with 
compound leaves (Maxonia). It exists in taxa that are terrestrial, taxa that are 
epiphytic and taxa that are aquatic. It is found in taxa with upright stems, with 
creeping stems, and with climbing stems. Xerophytes and hydrophytes, arctic and 
tropical ferns —— all have dimorphic representatives, the common species as well 
as the rare. 


260 Gardens’ Bulletin, Singapore — XXX (1977) 


For what ecological reasons has dimorphic evolution occurred in such a 
gamut of ferns? One can, of course, ask the same question regarding other 
pteridophytes ——- Equisetopsida and Lycopodiopsida which have made 
analogous changes of shoots rather than leaves. In all likelihood, similar forces have 
operated to produce strobili (analogous to apically hemidimorphous fronds) and 
whole fertile shoots (analogous to fertile-sterile segregation of whole fronds, for 
example Equisetum arvense L. and Lycopodium carolinianum L.). 


Some major extrinsic factors to take into account are seasonality, radiant 
energy (e.g., heat, light), role of air and water, and biotic factors (e.g., shading, 
grazers, fungi). The plant functions of most importance here are photosynthesis, 
spore production, and spore dispersal and establishment. 


All of the following adaptations may contribute to the differentiation of sterile and 
fertile fronds: 


1. To elevate spores of the fertile fronds into the wind stream for maximum 
dispersal. 


2. To permit more extreme drying effects to enable sporangia to open and 
close efficiently. . 


3. To spread out photosynthetic blades in horizontal positions for maximum 
reception of light. 


4. To place photosynthetic blades in most humid surroundings to counteract 
water loss. 


5. To bring photosynthetic blades closer to the substrate where there are 
higher carbon dioxide concentrations during day time. 


6. To enable differentiation of seasonal timing for most favourable period for 
spore reproduction and most favourable period for vegetative assimilation. 


7. To separate fertile from sterile parts if one or the other is more liable to 
grazing or fungal attack. 


Most authors agree that sporophylls tend to bring the spores into a position 
where they can better be carried away by the wind. Holttum (1938) describes the 
common situation which ‘takes the form of a somewhat contracted lamina in the 
fertile fronds, which stand erect, on longer stipes than the broader sterile fronds 
which bend away in a rosette around them, ... This form of dimorphism un- 
doubtedly helps the distribution of spores, raising the sporangia well above the 
surrounding leaves and exposing them to drier air and any slight wind that may 
exist.”” 


Copeland (1906) emphasized the drying aspects in his analysis. In his words, 
‘“‘Dimorphism, whether merely begun, or highly developed, whether a character of 
whole fronds or of their parts, has in all cases the object of permitting the proper 
dryness of the mature sporangia without an improper desiccation of the vegetative 
structures, This is done by merely raising the reproductive structures farther above 
the substratum; or (rarely) by special structural devices, such as notched margins; 
by a restriction of the assimilating surface of the reproductive frond or region, so 
that it may be sacrificed in emergency; or by a more complete elimination of the 
vegetative structures in constitutionally ephemeral fertile fronds.” 


_ As already stressed, origin of sporophylls is accompanied often by increasing 
divergence or specialization of the trophophylls or trophophyllary blade parts. This 
Specialization invokes progressive broadening of the blade together with re- 
orientation into spreading or horizontal position. The blade becomes shorter and 
if the fronds are clustered a rosette is formed (Fig. 2, D). In long-creeping 
thizomes, the much reduced trophophylls are scattered and often cling to the 
substrate. The modification of the trophophylls provides maximum photosynthetic 
surface for the area of the blades and orients them so that they have a moister, 


Leaf Dimorphy in Ferns 261 


more protected environment. Under some circumstances, greater assimilation of 
carbon dioxide may be possible due to diffusion from the substrate of the respira- 
tory products of bacteria, fungi, animals, and roots. 


We need to know more about optimal times of appearance of sporophylls and 
trophophylls. Time of spore production, for example, may be highly adaptive, but 
we will not know this until we understand more about life cycles in the natural 
environment. In the Great Lakes area of North America, Hill and W. Wagner 
(1974) determined that thin-walled green spores are discharged in the spring and 
very thick-walled, non-green spores are discharged in the fall for the most part. 
This may have to do with germination and establishment, as the former group of 
spores are adapted for quick germination on mineral medium and the latter for 
slow germination on organic medium. The best season for release of thin-walled 
spores may be the spring when the greatest moisture and rainfall are available. 
Comparable conditions prevail in the tropics with respect to wet and dry seasons. 


Drought conditions in tropical dry seasons may call forth extreme adaptations 
of the trophophylls, exaggerating those described above. as found in Anemia 
Sect. Trochopteris (A. elegans (Gardn.) Presl, A. eximia Taubert) with extremely 
small, flat laminae (cf. Mickel, 1962). The polypod, Microgramma heterophylla 
(L.) Wherry: illustrates an elaborate form of heterophylly in a vining fern. This 
tropical American fern grows over dry limestone rocks and produces short, clasping, 
ovate leaves with blunt tips, in contrast to a more upright, elongate form of sterile 
leaf, and the long, erect, narrowly lanceolate fertile leaves. The completely fiat. 
rosette-like sterile segment of the winter grapefern, Botrychium lunarioides, is 
probably adapted to growth only during winter months and is a device to capture 
maximum heat and light as well as to protect the tissues from sudden drying or 
cold shocks. 


Whether or not sporophyll-trophophyll divergence has adaptive value with 
respect to parasites is presently purely speculative, but such a possibility should 
not be ignored. Collectors notice frequently, especially in the tropics, that soral 
structures seem to be especially liable to molds and to tiny insects that feed 
upon the spores and sporangia. Conceivably there are adaptive advantages to 
isolating the parts most likely to be eaten from those which photosynthesize. There 
is also a chance that the eating of sori and fertile pinnae by small animals may 
play some role in dispersal, especially of those spores that germinate beneath the 
soil or under leaf litter. 


Although the onocleoid ferns are commonly cited by morphologists to 
illustrate foliar dimorphy, they are, in fact, unusual and not typical in several 
respects. Lloyd (1971) in his recent monograph of the systematics of the onocleoid 
ferns compared the fertile and sterile fronds of the three genera involved, but he 
did not compare them with other fern genera. Too little is known as yet about the 
rare Mexican genus Onocleopsis in its natural setting to evaluate it, but the two 
other genera, Onoclea and Matteuccia, show a number of unique features: The 
fertile leaves are (1) shorter than the sterile or equal to them, (2) their tissues are 
more dense and sclerified, (3) the sori, which are indusiate, are tightly encapsulated 
at maturity, and (4) the sporophylls persist much longer than the sterile fronds, 
which are early-deciduous in the autumn. What governs the final opening up of 
the bead-like pinnules is unknown, but in the eastern United States and Canada, 
where there is a strong winter-summer seasonality, the fertile units remain rigidly 
closed during the non-growing season and spore discharge is prevented. The hard, 
upright fertile leaves persist through the winter, covered with, or protruding 
through, the snow. The spores within stay green during their winter dormancy, and 
when they are released early in the spring they are ready to germinate immediately, 
thus taking advantage of the vernal moistness (Hill and W. Wagner, 1974). 
Although the sporophylls of Onoclea may sometimes equal or even surpass the 


262 Gardens’ Bulletin, Singapore — XXX (1977) 


trophophylls in height, those of Matteuccia are relatively much shorter, one-third or 
less the length of the trophophylls. In August and September, the sporophylls of 
Matteuccia are hard to see, buried as they are among the tall trophophylls and 
the other foliage of their swamp habitats. But in mid-winter all or most of the 
herbaceous plants have wilted and collapsed, so that the sporophylls are entirely 
exposed and easily swept by the wind. However, the actual discharge does not 
occur until March and April. Obviously the morphological peculiarities of these 
plants are correlated with the unusual seasonality of their spore production and 
release. 


DISCUSSION 


The noted Cornell University morphologist of the first half of this century, 
Arthur J. Eames, entertained some unusual theories about foliar dimorphism, In 
general he read the sequence just the opposite of the traditional interpretation and 
the one adopted here. In Eames’ words (1938), ““The segregation of vegetative and 
reproductive functions in different parts of the leaf, or in separate leaves, represents 
undoubtedly a primitive condition ...; from this ancient dimorphism there has 
developed the condition often called monomorphism -~—— found in the 
majority of ferns, a blade serving for both vegetative and reproductive functions 
... And from this there has developed a new dimorphic condition fertile and 
sterile leaves separate; this change has taken place independently in various genera 
...”’ He thus postulated “‘ancient” dimorphism, which is the primitive state, leading 
to monomorphism, and then, once again to “‘modern’”’ dimorphism. (Cf. his 
figure 175). 


Eames cited three families Ophioglossaceae, Osmundaceae, and 
Schizaeaceae as representing the primitive type. All of these are of the type 
referred to in this paper as having “‘botryoidal” sporangial arrangement. Those of 
‘“‘Polypodiaceae”” (ie., all higher leptosporangiate ferns) represent Eames’ 
‘“‘modern” type. We are not clear, however, upon what evidence he based his 
conclusions. The Ophioglossaceae are so isolated and so lacking in fossil evidence 
that we can only reason about what its course of evolution has been. We assume 
that it was derived from monomorphic fern-like plants because of analogies with 
repeated evolutionary sequences that we know in other groups. With respect to 
Osmundaceae, Miller (1971) concludes that “‘isomorphism represents the primitive 
state, incomplete dimorphism (only certain pinnae of a fertile frond modified for 
sporangia formation) is intermediate, and complete dimorphism is advanced,” and 
he calls attention to the fact that fragments of laminate fronds bearing osmunda- 
ceous sporangia occur mainly in Jurassic sediments but range throughout the 
Mesozoic. Miller notes that “‘such fronds are generally included in the form genus 
Todites.”” Even though dimorphic pinnae in Osmundaceae were also found as early 
as the Jurassic and Triassic, this does not mean that this condition was primitive, 
any more than in families known primarily only today in which both monomorphic 
and dimorphic pinnae occur. 


Also, as regards Schizaeaceae, the investigations of Mickel (1962, 1967) tend 
to contradict the conclusions of Eames. Mickel states that “‘although the change 
to dimorphism seems spectacular, in Anemia it is apparently a relatively simple 
and taxonomically minor one, which has occurred at least twice in the genus.” He 
explains the unique leaf structure of typical anemias with their upright fertile basal 
pinnae to be derived from the typical fern leaf with unmodified basal pinnae (as 
in the closely related genus Mohria), and he describes a series of species, beginning 
with Anemia colimensis Mickel, with nearly monomorphic pinnae, and continuing 
through more and more differentiated forms, that finally terminates in such end- 
points as A. rutifolia and A. millefolia. 


In view of the patterns seen in Osmundaceae and Anemioideae, and the total 
pattern of ferns in general, it is unlikely that there are two kinds of dimorphism, 


Leaf Dimorphy in Ferns 263 
‘primitive’ and ‘‘modern,” and that the latter arose from the former via a 
monomorphic stage. Such an hypothesis appears overly complex in the face of the 
over-all evidence. The hypothesis of one-way directionality is more economical and 
more probable. And when one considers the numerous analogous trends as are 
observed in other vascular plant orders such as Lycopodiales, Selaginellales, 
Equisetales and Cycadales, for example, the idea of reversal from dimorphic to 
monomorphic seems even less likely. 


In his rather lengthy discussion of foliar dimorphy, Dickason (1946) wrote 
that “‘Evidently dimorphism has not arisen in response to any given set of environ- 
mental factors, and cannot be considered as adaptive.” He based his conclusion, 
apparently, on the fact that dimorphy appears in plants of such widely different 
habits and habitats. Dickason was also concerned with the fact that “dimorphic 
and monomorphic species grow side by side.”” These conclusions are in keeping 
with a general negativism on Dickason’s part with respect to adaptive values of 
most phylogenetic trends in ferns. He did concede, however, that dimorphy is an 
evolutionary advancement over monomorphy. 


Whether Dickason’s conclusion that there is no adaptive value in dimorphy 
is justified is questionable. Merely that foliar dimorphy occurs in a wide spectrum 
of habits and habitats does not necessarily militate against its being adaptive with 
respect to one or more of the factors outlined above. Furthermore, examples of 
non-adaptive and adaptive character-states in taxa that grow side-by-side are 
numerous in all plant communities. The point is that all plant species probably 
have mixtures of both adaptive and non-adaptive or neutral attributes. 


Taxonomically there is no question that dimorphism is an extremely helpful 
character for making identifications. In Genera filicum, Copeland (1947) uses 
fertile-sterile dimorphism as a key character for over fifty genera. Nevertheless, it 
is not necessarily a valid character for establishing affinities or delimiting genera. 
For one thing, dimorphy is not a single character but an ensemble of different 
characters (Table 1), and any given expression invokes its own combination. Each 
manifestation must be analyzed accordingly. Furthermore, dimorphy has arisen in 
practically all families of ferns and in some of them probably several times. Table 
2 summarizes the incidence in the various families and subfamilies of ferns. What 
is designated as ““weak”’ involves only a few representatives of the taxon or only 
slight subdimorphism or both. “Strong” involves a majority of members and 
usually very distinctive sporophylls and trophophylls or well marked hemi- 
dimorphism. At the family level, dimorphy seems to be most pervasive in 
Ophioglossaceae, Plagiogyriaceae, Cheiropleuriaceae, Marsileaceae, and Salvinia- 
ceae. 


In spite of the fact that dimorphy has been used to distinguish a number of 
fern genera, it is questionable whether it is of fundamental importance. Sometimes 
the dimorphic taxon is connected by intermediates to monomorphic taxa. Tagawa 
and Iwatsuki (Iwatsuki, 1961) erected the genus Dimorphopteris on the basis of its 
perfectly dimorphic fronds, the fertile pinnae so contracted as to be linearimonili- 
form. However, the type species, D. moniliformis from the island of Halmahera in 
the Moluccas has been shown by Holttum (1972) to be an extreme in a series of 
subdimorphic members of a section of the genus Pronephrium. Copeland, among 
others, paid considerable attention to sporophyll-trophophyll distinction in his 
classification, but this was apparently partially a result of his emphasis on con- 
venience. In Table 3 are listed 12 genera (right-hand column) recognized by Cope- 
land which may indeed be too little differentiated to justify separation from their 
parent genera. 4 


4]It may have some relevance that two recent students of the genus Equisetum, namely R. L. 
Hauke and C. N. Page, have rejected shoot dimorphism as a determining character for 
establishing sections and subgenera (Hauke, 1974). 


264 Gardens’ Bulletin, Singapore — XXX (1977) 


The major taxonomic value of foliar dimorphy in ferns as a whole is at the 
species level. There are numerous examples of pairs of closely related species in 
which one member is essentially monomorphic and the other dimorphic, Such 
pairs deserve close examination; because of their genetic similarity their study 
may give us valuable insights into the evolutionary and adaptive significance of 
this phenomenon. 


Table 1. Parallel and convergent changes associated with sporophyll-trophophyll differentia- 


tion. The primitive state for each character is undifferentiated or nearly so. 


LEAF SIZE 


ORIENTATION 
PETIOLE LENGTH 
BLADE OUTLINE 
BLADE CUTTING 


LAMINAR WEBBING 


LAMINAR MARGINS 


VENATION COMPLEXITY 


VEIN INTERVAL 


MESOPHYLL 


EPIDERMIS 


SORIATION 


ENCAPSULATION 


SEASONALITY 


DURATION 


Strongly unequal 

Sporophyll erect, trophophyll spreading 
Unequal 

Sporophyll narrow, trophophyll wide 


Sporophyll usually more divided; trophophyll 


usually less divided (but exceptions) 


Sporophyll with reduction or loss of lamina; 
trophophyll unchanged or with increase 


Sporophyll commonly smoother; trophophyll 
more toothed 


Sporophyll most commonly simplified by vein 
loss; trophophyll more complex 


Sporophyll veins more remote; trophophyll 
veins more approximate 


Sporophyll non-green, soft; trophophyll green, 
more compact and rigid 


Sporophyll with fewer stomates and _ larger, 
smoother-walled cells; trophophyll with 
normal epidermis 


Sori concentrated, often confluent into acrosti- 
choid condition or grouped into botryoidal 
condition 


Sporangia, at least while young, covered by 
rolled lamina 


Sporophyll separated from trophophyll in time 
of appearance, usually earlier but may be 
later 


Sporophyll short-lived; trophophyll long-lived. 
Rarely otherwise 


Leaf Dimorphy in Ferns 265 


Table 2. Parallel and convergent evolution of foliar dimorphy in ferns. Relative incidence 
and main types in different families or subfamilies. Letters refer to dimorphic types in 
Figure 1. 


STRONG TO 


MODERATE COMPLETE 


ABSENT TO WEAK 


Polypodioideae (B, D) Ophioglossaceae (C)} 


Marattiaceae (D) 


Gleicheniaceae Lygodioideae (B) Osmundaceae (B, D, E) 
Matoniaceae Dryopteridoideae (B,D, F) Plagiogyriaceae (D) 
Dipteridaceae Blechnoideae (D) Anemioideae (C)? 
Loxogrammeoideae Ceratopteridoideae (D) 


Marsileaceae (C)4 
Salviniaceae (D)® 


Grammitoideae (B, D) 
Schizaeoideae (D)?* 
Adiantoideae (D) 
Vittarioideae 
Cyatheoideae 
Hymenophylloideae 
Lindsaeoideae 
Asplenioideae 
Azollaceae 


1 Fertile spike composed of fused basal fertile pinnae. 

2 Fertile basal pinnae not fused, except in some cases to the petiole. 

3 Fertile blade drastically reduced and petiole exaggerated, Sterile petioles maintained at 
maturity in Schizaea pusilla Pursh. 

4 Type “C” if sporocarps are modified fertile pinnae. 

5 Sporophyll is the immersed root-like leaf. 


Table 3. Genera of close affinity which were discriminated by Copeland (1947) on the 
basis of fertile-sterile leaf dimorphism. 


MONOMORPHIC OR 
SUBDIMORPHIC 
PARENTAL GENUS 


DIMORPHIC SEGREGATE 


POLYPODIACEAE 
Pleopeltis Lemmaphyllum 
Pleopeltis Marginariopsis 
Colysis Dendroglossa 
Aglaomorpha Merinthosorus 
Crypsinus Pycnoloma 
ADIANTACEAE S. l. 
Pteris Neurocallis 
CYATHEACEAE S. 1. 
Trichomanes Feea 
ASPLENIACEAE S. 1, 
Phanerophlebia Cyclodium 
Ctenitis Psomiocarpa 
Tectaria Tectaridium 
Tectaria Hemigramma 
Tectaria Quercifilix 


266 Gardens’ Bulletin, Singapore — XXX (1977} 


LITERATURE CITED 
Bower, F. O. 1923. The Ferns. Vol. I. Analytical Examination of the Criteria of 
Comparison. Cambridge U. Press. 


Bower, F. O. 1928. The Ferns. Vol. III. The Leptosporangiate Ferns. Cambridge 
WaPress, 


Copeland, Edwin B. 1907. The comparative ecology of San Ramon Polypodiaceae. 
Philippine Jour. Sci. 2 (no. 1): 1-76, pl. 1-4. 


Copeland, Edwin B. 1947. Genera filicum. Chronica Botanica, Waltham, Mass. 


Dickason, Frederick Garrett. 1946. A phylogenetic study of the ferns of Burma. 
Ohio Jour. Sci. 156 (no, 2): 73-108. 


Eames, Arthur J. 1936. Morphology of Vascular Plants. Lower Groups. McGraw- 
Hill Co., New York. 


Gupta, K. M. and T. N. Bhardwaja. 1956. Indian Marsileas: their morphology 
and systematics. Jour. Bombay Nat. Hist. Soc. 53: 423-444, pl. 1-3. 


Gupta, K. M. 1962. Marsilea. Botanical Monograph No. 2 Council of Sci. and Ind. 
Res., New Delhi. 


Hauke, Richard L. 1974. The taxonomy of Equisetum ——— an overview. New 
Botanist 1: 89-95. 


Hill, Royce C. and W. H. Wagner, Jr. 1974. Seasonality and spore type of the 
pteridophytes of Michigan. Mich Bot. 13: 40-44. 


Holttum, R. E. 1938. The ecology of tropical pteridophytes. Jn Verdoorn, F., 
Manual of Pteridology, pp. 420-450. Martinus Nijhoff, The Hague. 


Holttum, R. E. 1939. The genus Lomariopsis in Madagascar and the Mascarene 
Islands. Notul. Syst. 8: 48-62. 


Holttum, R. E. 1972. Studies in the family Thelypteridaceae. The genus Proneph- 
rium Pres]. Blumea 20: 105-126. 


Iwatsuki, Kunio. 1961. Taxonomic studies of Pteridophyta VI. Acta Phytotax. 
Geobot. 19: 1-11. 


Labouriau, Luiz Gouvéa. 1958, Studies on the initiation of sporangia in ferns. 
Arquivos do Mus. Nacional, Rio de Janeiro 46: 119-202. 


Lommasson, Robert C. and C. H. Young, Jr. 1971. Vascularization of fern leaves. 
Amer. Fern Jour. 61: 87-93. 


Lloyd, Robert M. 1971. Systematics of the Onocleoid ferns. Univ. Calif. Publ. Bot. 
61: 1-93. | 

Mickel, John T. 1962. A monographic study of the fern genus Anemia, Subgenus 
Coptophyllum. Iowa State Jour. Sci. 36: 349-482. 


Mickel, John T. 1967. The phylogenetic position of Anemia colimensis. Amer. Jour. 
Bot. 54: 432-437. 


Nozu, Yoshitomo, 1968. Studies on the leaves of the Ferns. I. The dissimilarities 
of the fertile and sterile fronds in some ferns. Jour. Fac. Sci., Univ. Tokyo 
Sec. III. 10: 13-27, pl. 2. 


Wagner, W. H. Jr. 1965. Paraphyses: Comments on recent papers. Taxon 14: 
299-302. | 


Leaf Dimorphy in Ferns 267 


Wagner, W. H., Jr. 1973. Some future challenges of fern systematics and phylogeny. 
In Jermy, A. C., Crabbe, J. A., and Thomas, B. A. (ed.), ““The Phylogeny and 
Classification of the Ferns.’ Bot. Jour. Linnaean Soc. 67 (Suppl. No. 1): 
245-256. 


Wagner, W. H., Jr., D. R. Farrar, and B. W. McAlpin. 1970. Pteridology of the 
Highlands Biological Station Area, Southern Appalachians. Jour. Elisha 
Mitchell Sci. Soc. 86: 1-27. 


White, Richard A. 1971. Experimental and developmental studies of the fern 
sporophyte. Bot. Rev. 37: 509-540. 


Wieffering, J. H. 1964. A preliminary revision of the Indo-Pacific species of 
Ophioglossum (Ophioglossaceae). Blumea 12: 321-337. 


why. -: 4 
J J 


- 


+. F907: The Rants 
‘ ily yn" 


nal idee * asibute” comet vf i 
ws L929 | ir RODE a ‘ naa { +0 
+ ? tion ‘Oba: nl ) , 
lo Vo) 14 ‘ | 


be 


y garnuls 


larsileasr 
oa 


A Note on Dendrobium serpens (Hk.f.) Hk.f. 
by 


GUNNAR SEIDENFADEN 
Botanical Museum, Copenhagen 


In his “Orchids of Malaya’, Holttum (1964 : 319) lists Dendrobium serpens, 
adding to his description that “‘more information is wanted about this species”. 
We listed it in “Orchids of Thailand’? (Seidenfaden & Smitinand 1960: 244) 
based on Holttum’s information that Dendrobium virescens Ridl. should be con- 
sidered a synonym, and remarked that the plant was found once in Tomoh in 
Peninsular Thailand just north of the Malayan border but we had never ourselves 
seen the plant. 


In the summer of 1970 Smitinand and I went on an orchid hunting expedition 
to the southernmost provinces of Thailand. Among the plants collected near the 
border of Malaya at Waeng Forest Station west of Sungai Kolok was a trivial 
Dendrobium, which was brought to the hothouses in Copenhagen in the hope that 
it would produce the flowers necessary for the identification. No flowers appeared 
for years, but when in the spring of 1975 we experimentally raised the average 
humidity in the houses, several of the plants showed increased activity, and, in 
May 1975 it produced flowers, making it possible to identify it as Dendrobium 
serpens by means of Holttum’s keys. This led to the small study related below, 
giving some bits of the information wanted by Holttum. 


During the period of printing of the orchid part of ““The Flora of British 
India” Hooker f. continued to receive new material, both in the form of new 
collections in the field and as drawings, mainly from the Calcutta herbarium. This 
necessitated the addition of a supplement to his treatment, found in the sixth 
volume of “Flora of British India’? containing a lot of important, although too 
often overlooked, information. 


Among the drawings received from Calcutta was one of a Dendrobium of the 
Pedilonum section, said to represent a plant growing at ‘“‘Rangoon’’, brought by 
Mr. Gilbert, a resident of Moulmein, where he succeeded Parish, and on a more 
modest scale tried to take up the work of this great botanist, sending to Europe 
herbarium specimens of orchids, often accompanied by sketches of which several 
are found in Reichenbach’s herbarium in Vienna. In the present case Gilbert must 
have sent a living plant, because when Hooker f. received the drawing from 
Calcutta, to it was attached a single dried flower, said to have developed in the 
Garden at Calcutta in November 1884. As to my knowledge the species has never 
turned up in the Pegu area, I feel we must not exclude the possibility that the plant 
was not at all found “at Rangoon” but rather, brought up to Rangoon from 
Moulmein for shipment to Calcutta. Hooker f. called the plant D. panduriferum. 


Another drawing received from Calcutta, made after a plant collected by 
Kunstler in Perak and flowering in Calcutta in May 1884, prompted Hooker f. 
to describe it as a variety of Gilbert’s plant, calling it Dendrobium panduriferum 
var. serpens. Of this he evidently got no plant material and had in his description 
to rely solely on the drawing. This too he later reproduced (1895:10, PI.16), at 
which time he reached the opinion that the two entities were different enough to 
justify considering the Perak plant a separate species, Dendrobium serpens. 

Very little information on these two taxa has appeared since. Grant (1895: 108) 
copied Hooker f. in his Burma list, while Ridley in his paper on the orchids of 


269 


270 Gardens’ Bulletin, Singapore — XXX (1977) 


GT 7517 


RIDLEY 14303 


MACHADO 


Fig. 1. Dendrobium serpens (Hk. f.) Hk. f. a. inflorescence from a fresh Thai specimen, 
b. flower from the Temengo specimen, c. flower from the type specimen of D. virescens, d. 
epichile and e. flower without lip and mentum from the Thai specimen. 


Dendrobium ser pens 27] 


the Malay Peninsula (1896) overlooked Kunstler’s Perak plant which he sub- 
sequently listed in his “‘Materials” (1907:47), and, in his “‘Flora of the Malay 
Peninsula” (1924:47) we learn that he had identified one of his own collections 
as being Kunstler’s plant, giving the locality as Ulu Temengo (= Temengor) on 
an island in the Kertai River. 


Already in 1896 Ridley had, however, introduced a new taxon, Dendrobium 
virescens, based on a collection made by Machado further to the Northwest, at 
Tomo (spelled by Ridley as Tomoh) which I believe is on the Thai side of the border 
and, in the “‘Materials’’ we learn that he had identified a plant collected at Taiping 
in Perak by Curtis, — the latter is repeated in the 1924-paper. 


In his large monograph on Dendrobium, Kranzlin (1910) lists Dendrobium 
virescens and D. panduriferum without adding any new information, he places 
them in two different subsections of the Pedilonum-section, although they might 
be considered conspecific, cfr. below. Dendrobium serpens he choses to consider 
a synonym of Reichenbach’s Dendrobium ionopus, a plant said by Reichenbach to 
have come from Burma. Hooker f. had originally suggested that his var. serpens 
could be the same as D. ionopus, but later (1895:11) he indicated that he was 
convinced of the contrary; Kranzlin seems to disregard this viewpoint, while both 
Ridley and Holttum ignore Kranzlin’s opinion. 


In response to “more information” sought by Holttum, I got with the kind 
assistance of Dr. Chang the available material from Singapore, at the same time 
receiving from Vienna the material kept in the Reichenbach herbarium under the 
name of Dendrobium ionopus. 


There seems no doubt that the recently found Thai plant is identical with both 
the Tomo plant which is the type specimen of Dendrobium virescens Ridl. and 
with the plants from Ulu Temengo and Taiping. The most characteristic features 
are details of the lip, see Fig. 1. There is a half-moon shaped transverse callus at 
the base of the epichile at the side of which the edges fold into ears continuing 
as longitudinal somewhat converging folds on the blade of the epichile. There is 
no fattening on the epichile, the whole of it is of very thin texture, which is not very 
clear on the drawings Hooker f. received from Calcutta; based only on these, 
wane ails the blade as “‘cordate above and below, slightly constricted at 
the middle”’. 


When comparing the original drawing which was the only basis for Hooker’s 
analysis, with the plate published by Hooker of Dendrobium panduriferum it is 
important to recall that here the details of the flower are based on Hooker’s own 
dissection of a flower he received with the habit drawing. It is tempting to suggest 
that after moistening or cooking the flower, any folds and ears in the thin material 
would vanish, and a lip like the one found in the material we have from Thailand 
and Malaya of D. serpens would look pretty much like Hooker’s drawing of the 
lip of Dendrobium panduriferum, described by Hooker as “‘deeply constricted with 
auricle-like sidelobes”, the transversal callus is the same. I am therefore quite 
convinced that D. panduriferum and D. serpens are the same species. The only 
depiction of a flower in front view in the habit drawing of D. panduriferum (drawn 
in India after fresh material) shows, in my opinion, an epichile indistinguishable 
from the one shown in the Indian picture of D. serpens, and the other differences 
mentioned by Hooker seem not very convincing. I feel, however, that it would be 
too hasty to reduce D. serpens to a synonym, perhaps it is safer to wait until fresh 
material has been obtained of plants from ‘‘Rangoon’’; repeating Holttum’s words 
that “‘more information is wanted’’, my preliminary summary on D. serpens 
therefore reads as follows: 


Dendrobium serpens (Hk. f.) Hk. f., Ann. Bot. Gard. Calc. 5: 10, T. 16, 1895.— 
ee 1907:51; idem 1924:47; Holttum 1964: 319; Seidenfaden & Smitinand 
1960: 244. 


272 Gardens’ Bulletin, Singapore — XXX (1977) 


Dendrobium panduriferum var, serpens Hk. f., Fl. Brit. Ind. 6: 186, 1890. 


Dendrobium virescens Ridl., Journ. Linn. Soc. 32:259, 1896—F. N. Williams 
1904: 368; Ridley 1907:51; Kranzlin 1910:100; Ridley 1924:47. 


Dendrobium ionopus auct. non Rchb, f.: Kranzlin 1910: 122 (p.p.) 


Distribution: Thailand: Legeh at Tomoh (Macado s.n. SING!, type of Dendrobium 
virescens); Near Waeng Forest Station, Narathiwat 250m (GT 7517 Cl). 


Malaya: Perak (drawing in Calcutta after a plant sent by Kunstler, type); Ulu 
Temengo on an island in Kertai River (Ridley 13303 SING!); Taiping (Curtis 
s.n. SING)!); presumably also Karangan, Kedah (Haniff s.n. SING!, insufficient 
material). 


Now, on Dendrobium ionopus Rchb, f. of which Kranzlin considered Dendro- 
bium serpens a synonym: 


The material of this taxon in Herb. Reichenbach consists of two sheets. The 
one carrying the handwritten original description (No. 41205) contains a single 
dissected flower which, under the number 3769, had been sent by Low in November 
1882 and said to originate in Burma, further, some rough coloured sketches by 
Reichenbach of that flower, and a copy of one of Day’s drawings. I later saw the 
original of that drawing in Day’s rich manuscript collection in Kew, made in 
November 1882, and a note gives the same information that it was imported by 
Mr. Low from Burma. The other sheet (No. 41204) contains an inflorescence also 
sent by Low, with whom it flowered in 1881; it is noted that the collector was 
Boxall and that it originated in the Philippines. 


The flowers on the two sheets are identical. The most important characteristics 
are the two low triangular lateral calli near the edge at the base of the epichile 
(these are both in Reichenbach’s and Day’s drawings coloured dark red-purple), 
and the obtuse tooth in the “‘spur’’, see Fig. 2. The flower is quite different from 
that of D. serpens, and Kranzlin was wrong. 


But seeing my sketches of D. ionopus, Dr. Garay called my attention to the 
figure of a taxon described in 1908 by Kranzlin as a new species, Dendrobium 
epidendropsis, based on one of Loher’s plants. This is clearly the same as Low’s 
plants. D. epidendropsis has later been recorded several times from Luzon. 


Here again, it is at hand to be suspicious about the old information. Con- 
sidering the general tendency during the later part of the last Century to be less 
exact (and often with purpose misleading) on the origin of commercially introduced 
plants, I feel convinced that the flower Low sent to Reichenbach and Day in 1882 
as “from Burma” is from the same plant or the same consignment as the one sent 
a year before as being a Boxall-collection from the Philippines, and I feel we 
should exclude the record from Burma. 

Finally, I would suggest that the photograph reproduced under the name 
Dendrobium O’Brienianum by Davis & Steiner (1952: 120) looks more like 
Dendrobium ionopus; Kranzlin’s drawing of D. O’Brienianum (1910: Fig. 5 N-O) 
has a much more slender mentum, also the photograph shows a pair of dark spots 
somewhere at the base of the lip, not mentioned in Kranzlin’s colour description, 
but, naturally, the photo is not sharp enough for a sure identification. 
pee My knowledge on Dendrobium ionopus could therefore be summarized as 

ollows: 


Dendrobium ionopus Rchb. f., Gard. Chron. 1882, 2: 808.— 
Hooker f. 1890: 732, Grant 1895: 100; Kranzlin 1910: 122. (excl. syn. D. serpens). 


Dendrobium epidendropsis Krzl., Orchis 2: 79, Fig. 12, 1908, syn. nov. — Ames 
1909: 598; Kranzlin 1910: 106; Ames 1915: 123; idem 1925: 348. 


Dendrobium ser pens 273 


. “ ae 
2 


aa i 
Pan ee a8 
RR tt Ch 


5mm 


H.R. 41205 


Fig. 2. Dendrobium ionopus Rchb. f. Type specimen. a. the only mutilated flower, with 


dorsal sepal, a petal and part of a lateral sepal missing, lip removed, b. epichile of lip, c. 
tooth in mentum. 


Dendrobium O’Brienianum auct. non Kranzlin: Davis & Steiner 1952: 120? 


Distribution:? Burma: s.n. (Low 3769 Herb, Reichenbach 41205! 
improbable). 


, type, origin 
Philippines: sine loc. (Boxall s.n. Herb. Reichenbach 41204!; Loher s.n., type of 
D. epidendropsis; Lyon 118 and 161, fide Ames); Lamao River, Bataan, Luzon 


(Borden 2109 and Elmer 6839, fide Ames); Rizal, Luzon (Ramos 3060, fide 
Ames). 


Literature Cited 


Ames, O., 1909: Notes on Philippine orchids with descriptions of new species 1. 
Phil. Journ. Sci. 4, 5: 593-600. 


. 1915: Orchidaceae 5. The Genera and Species of Philippine Orchids. 
Boston, 271 pp. 


. 1925: Enumeration of Philippine Apostasiaceae and Orchidaceae. Bur. 
Sci. Publ. No. 18: 252-463. Manila. 


Davis, R. S. & M. L. Steiner, 1952: Philippine Orchids, New York, 270 pp. 


274 Gardens’ Bulletin, Singapore — XXX (1977) 


Grant, B., 1895: The Orchids of Burma. Rangoon, 416 pp. 
Holttum, R. E. 1964: Orchids of Malaya. Flora of Malaya 1. 3rd. ed. Singapore. 


Hooker, J. D., 1890: The Flora of British India 5: 667-864, 6: 1-198. Ashford, 
Kent. 


. 1895: A Century of Indian Orchids. — Ann, Roy Bot. Gard. Calc. 5: 
1-68, Pls. 


Krinzlin, F., 1910: Orchidaceae-Monandrae-Dendrobiinae, in Engler: Das 
Pflanzenreich IV. 50. II. B. 21. 


Ridley, H. N., 1896: The Orchideae and Apostasiaceae of the Malay Peninsula. 
Journ, Linn, Soc. Bot. 32: 213-416. 


. 1907: Materials for a Flora of the Malayan Peninsula 1: 7-233. 
Singapore. 


. 1924: The Flora of the Malay Peninsula 4: 4-233. Ashford Kent. 
Seidenfaden, G. & T, Smitinand: The Orchids of Thailand 2, 2: 185-326 Bangkok. 


Williams, F, N., 1904: Liste des Plantes connues du Siam. Bull. Herb. Boiss. 
2. ser. 4: 362-372. 


Notes on the Systematy of Malayan Phanerogams XXV.* 


Araliaceae 
by 
BENJAMIN C. STONE 


Department of Botany, University of Malaya, 
Kuala Lumpur 


Abstract 


Two genera are reduced to synonymy, Wardenia King to Brassaiopsis Decne. & Planch.. 
and Acanthophora Merr. to Aralia L. Critical notes on some other genera, some new 
combinations, and two new species, Brassaiopsis minor and Schefflera singularis, are published. 
A brief growth analysis of Arthrophyllum diversifolium sensu King is presented. 


In the course of preparing the treatment of Araliaceae for Vol. 3 of the “Tree 
Flora of Malaya” a number of problems involving not only the recognition and 
delimitation of species, but also of genera, have arisen. Two undescribed species 
have been encountered and are published here. Equally important, the status of 
two hitherto monotypic genera, Wardenia King and Acanthophora Merr., has been 
reassessed, and it has been concluded that they should be reduced to synonymy 
under Brassaiopsis Decne. & Planch. and Aralia L. respectively. The largest genus 
of the family in Malaya, Schefflera J. R. & G. Forst., is currently being revised 
by D. G. Frodin of the University of Papua New Guinea. In his dissertation 
(Cambridge University 1971) he has taken a broad view of Schefflera, incorporating 
within it three genera recognized by Ridley in his Flora of the Malay Peninsula 
(vol. 1, 1922), Brassaia Endl., Scheffleropsis Ridl., and Tupidanthus Hk. f,. & 
Thoms. In a recent publication (1970) Philipson has reviewed the genus Gastonia 
and remarked on a species which occurs in Malaya, which was not in Ridley’s 
Flora. The genus Acanthopanax Miq. has been found to occur in Malaya, the sole 
Malayan species being A. malayana Henderson (Gdns Bull. Sing. 7: 105-6, pl. 
23. 1933). Ridley himself described a second species of Hederopsis King some years 
after his Flora was published (H. major Ridl. in Kew Bull. 124. 1929). The genus 
Polyscias Forst. is not in Ridley’s Flora, but is represented in the Peninsula by 
several species found in cultivation. 

Thus at present there are known to occur in Malaya ten genera of Araliaceae: 
Acanthopanax, Aralia, Arthrophyllum, Brassaiopsis, Dendropanax, Gastonia, 
Hederopsis, Polyscias, Schefflera, and Trevesia. If Tupidanthus is maintained as a 
distinct genus, as advocated by Philipson (pers. comm.), the total is eleven. 

The genus Aralidium Mig. was assigned to Cornaceae in Ridley’s Flora, but 
placed in Araliaceae by Corner (Wayside Trees of Malaya 1: 154. 1940 & 1952). 
After considerable preliminary study it is deemed preferable to exclude it from 
Araliaceae on account of its lack of resin canals, among other characters. Whether 
it should rejoin Cornaceae or find some other position is a problem still under 
study. 


275 


276 Gardens’ Bulletin, Singapore — X XX (1977) 


Aralia L. 


Ridley reported three species of Aralia from the Malay Peninsula, A. thom- 
sonii Seem., A. armata (Wall. ex G. Don) Seem., and A. ferox Miq. Merrill 
(Philipp. J. Sci. 13, Cr, 316-318, 1918) showed that Philippine and Celebes 
specimens determined as A. ferox Mig. were not that species at all, and because 
of the climbing habit created a new genus Acanthophora for these plants with A. 
scandens Merr. as the sole species. Later, van Steenis (Bull. J. Bot. Buitenz. ser. 3, 
17: 390, 1948) added further determinations and included the “‘A. ferox” of 
Ridley’s flora, adding the genus Acanthophora to the known Malayan Araliaceae. 


The species Acanthophora scandens Mert. is certainly distinct. It has recently 
again been collected along the Ginting Highlands Road, Pahang (Stone 12016). 
A study of this material and a review of the pertinent literature suggest however 
that the generic status for this species is highly dubious, its sole claim to it being 
the lianoid habit. Since most species of Aralia are shrubs or trees (or herbs) the 
liane habit does stand out. On the other hand there are many genera known which 
include species with this habit along-side others which are trees or shrubs, witness 
Schefflera, and outside the Araliaceae, Derris (Leguminosae), to name but two 
examples. In this view I am supported by D. G. Frodin (Misc, Rec. Fl. Mal. 
Found. III: 8. 1973). 


In any case there is a nomenclatural reason why the name Acanthophora 
Merr. 1918 cannot be used, even if the generic concept it denotes were considered 
worth retaining. This is the generic name Acanthophora Lamouroux 1813, applied 
to a taxon of Rhodophyceae (Red Algae). This name was obviously overlooked 
by Merrill, when he created what was, in fact, a later homonym. 


Aralia scandens (Merr.) T. D. Ha, Novost. Sist, Vysh. Rast. 11: 229, 1974. 


Acanthophora scandens Merr. Philip. J. Sci. C, 13: 316-318. 1918. 
Aralia ferox sensu Ridley (non Miquel) in Fl, Mal. Pen. 1: 873. 1922. 
(Not Acanthophora Lamouroux, 1813 (Algae) ). 


Distrib. — Malaya, Philippines, Celebes. In Malaya: Perak, Pahang, and 
Selangor. Fig. 1. 


Arthrophyllum Blume 


Among the Araliaceae this is one of the more clearly distinct genera, rather 
easily recognized by its pinnate leaves and 1-celled ovary and fruit. The species 
are another matter. They are poorly described, mainly because of poor inadequate 
collections, and badly discriminated. Most are rather difficult to distinguish. Those 
in the Peninsula in particular are in a state of confusion because of the too- 
Optimistic classification by Ridley. In the Flora (I: 885) Ridley lists 8 species. 
These cannot be reached through the key and cannot be recognized from the 
descriptions which are too brief and contain errors of fact. The available specimens, 
including types and syntypes, even when examined, are not always decisive. Of 
the 8 species listed by Ridley, four seem to me reasonably clear; the others are 
to be considered as synonyms or as varieties of these four. 


The main point of this note is to describe in some detail the basic growth 
pattern of the largest of the Malayan Arthrophyllums. This plant was called A. 
diversifolium by Clarke, King and by Corner, while Ridley called it A. ovalifolium. 
Since it never possesses bipinnate leaves, it does not correspond very well with the 
former. Apparently it is not the same as the latter. The nomenclature will be treated 
In a separate paper by W. R..Philipson. At any rate, it is clear that throughout 
the lowlands of Malaya, there is a single, rather uniform tall Arthrophyllum species, . 


277 


Araliaceae 


‘yoriq ul uMOYsS ovuUId _JeTodyg,, ‘ainjonijs Jeo] 
JO WeISLIP 7Y8y ‘UOIDASIBUOT PUL ‘MOIA PUD ‘MOIA OPIS Ul Spoos—:d “}INIJ JO MOIA OPIS—: J “}IMIZ JO MOIA do}—:q “oyrsi—q 
‘9seq-YyouvIg YWM JWoUIsIs Wo}s—:d “(o1WUd9) soduULY Ivpndys suiddepioro YUM ‘opis roddn ‘Jeo, 1odunoA [[ys—:gq ‘Jeo, Surdoyaaop 
@unok pure ojndys pogo] Suimoys oseqjvsoy YM xode yooys—: VY ‘xode jooys jo siepp sifa7 ‘eH ‘A’ L C19) suapuvos vijosp 


SUPpHOIL DI PLYY Wogr 


—— 


oon mic 

! : ‘ | 

a4 | 
, 7 

hd 


SUaPUCIS EVE. 1 Wy 


| Bt 


278 Gardens’ Bulletin, Singapore — XXX (1977) 


which often enters waste ground as a pioneer regrowth plant. Its trunk eventually 
branches sparsely, and the trees may reach a height of over 15 m. This is the plant 
referred to here in the following description. 


Arthrophyllum sp. — A. diversifolium sensu King, Mat. Fl. Mal. Pen. in J. R. 
As. Soc. Bengal, 47, II (1): 59. 1898, non Blume; sensu Corner, Wayside Trees 
of Malaya 1: 155, 1952. — A. ovalifolium sensu Ridley, Fl. Mal. Pen. 1 : 885, 
1922, non Jungh. & De Vriese. Fig. 2. 


A. congestum Ridl. J. Fed. Mal. St. Mus. 10: 137. 1920; Flora 1: 887. 1922. 
Lectotype: SELANGOR: Klang Gates, Ridley 13421. (SING). 


Although Ridley’s binomial A. congestum applies unquestionably to this plant 
there seems little doubt that it is to fall into synonymy when a wider study of the 
group is made, and this is to be reported on by Professor Philipson in this volume. 


Ridley attributed the name A. ovalifolium to Miquel, but the authors, as 
stated by Miquel, were Junghuhn and De Vriese. Miquel’s description does not 
seem to be a different interpretation. In describing A. congestum Ridley notes the 
flowers as with 4 petals, but examination of the type collection shows that this is 
wrong; the flowers, as would normally be the case, are 5-merous. There are no 
other characters to distinguish this from the common, arboreous lowland species 
which Ridley calls A. ovalifolium. In accepting this name for a Malayan species, 
the chance of error is not small, and it remains to be checked against authentic 
specimens, At any rate, the species intended here is the most abundant of the 
Malayan lowlands, at least on the west coast, and is usually, when mature, a 
branched tree up to 14 m tall with a trunk reaching 30 cm diameter. The leaves 
on the main erect (orthotropic) stems are closely spiralled; they are large, once- 
pinnate, reaching a length of 2 meters, with 12-14 pairs of large leaflets (Fig. 2-2). 
Like Ridley, I have never seen a bipinnate leaf in any Arthrophyllum; on this 
basis I have refrained from identifying our plants with A. diversifolium Bl. or its 
synonyms, though this is the name used by King in the ‘Materials’ and Corner in 
“Wayside trees of Malaya (1940)” and found on a number of herbarium specimens 
in SING and KEP. According to the available descriptions A. diversifolium has at 
least some bipinnate leaves. It is possible that hasty observation of our plants would 
suggest that they too bear bipinnate leaves, but this arises through confusion of the 
axillary determinate branches (which bear the flowering shoots) with leaves. These 
axillary branches (Fig. 2-3) are more or less plagiotropic (though not usually 
horizontal), and bear at their widely spaced swollen nodes opposite bipinnate but 
reduced leaves, or nearer the tip, whorled and even further reduced leaves; in the 
axils of the latter arise the short flowering shoots (Fig. 2-3 and 2-9), which 
themselves have 1-3 nodes with pairs of simple leaves (Fig. 2-6). The resemblance 
of the lateral branch to the rachis of one of the big leaves is striking, but its 
position and smaller number of nodes serve to identify it. Since the leaves borne 
on the lateral branches are much smaller, and with successively reduced numbers 
of nodes, they are easy to collect, and hence are found on most herbarium sheets; 
but they are not really representative of the plants, as besides being smaller, they 
are not or scarcely asymmetric, which is the usual condition of the leaflets on the 
lower, big leaves. Also, their number of lateral nerves may be smaller. These 
Caeamaattes make identification of such fragmentary collections difficult and 

esitant. 


Plants of this species generally remain unbranched until they are about 
2.5 m tall; at this stage they may branch, but the branches are recapitulations of 
the trunk, i.e. orthotropic with spiral phyllotaxy and more or less equivalent. These 
may grow to an equal length, then flower. Flowering is by inception of the axillary 
lateral branches with distichous or whorled phyllotaxy, as described above; but 
new orthotropic shoots arise from other axils providing relays. The lateral shoots 
are strongly determinate in growth and reach about the same length as the big 


Araliaceae 279 


BCE. 12166 


i 


10 


q 


Wz 


N; 


\\) \\ F 
A 
NS ) () Y} 
ALG 
QV \ \ 4 aN 
~ O09 VYN 


Be i im y ( 
| ay, , 


» VO NUON. 


iors 


\ 
10 cm 2 


F.C. Stone 


ie) 
ofe/. 


6 


Arthrophyllum 


Fig. 2. Arthrophyllum sp. (A. diversifolium, sensu King). Aspects of its morphology. | :— 
First-order shoot. 2:—Pinnate leaf of first-order shoot. 3:—Second-order shoot. 
4:—Medial leaflet from leaf of first-order shoot. 5:—Medial leaflet from lowest 
leaf of second-order shoot. 6:—Simple leaf from third-order (flowering) shoot. 
7:—Cross-section of leaf rachis. 8:—Axillary bud. 9:—Apex of first-order shoot 
bearing second-order shoot branches (PB) and their axillant leaves (LF) or their 
scars (LS). 10:—Floral details. Stone 12156. 


280 Gardens’ Bulletin, Singapore — XXX (1977) 


pinnate leaves of vegetative stems. Their resemblance to rachises has been mentioned 
above; in fact, the immediate appearance suggests that the flowering umbels arise 
in the axils of leaflets, but close study dispels this impression. The big lower 
leaves always have at least 10 nodes, but the upper leaves on the lateral branches 
may have as few as 1 or 2 nodes (these leaves usually immediately subtend the 
flowering shoots). The reversion to spiral arrangement at the apex of the lateral 
branch is abrupt and striking. On the flowering shoots, there are umbels in the 
axils of the leaves on the median nodes as well as clustered at the apex. The 
inflorescence, therefore, consists, strictly speaking, of a pseudumbellate cluster of 
simple umbels, although superficially the appearance suggests a 2- or even 3-com- 
pound umbellate structure. 


The leaflets of a single leaf often have unequal laminal areas. The whole row 
of leaflets on one side (e.g. the “‘left’’ or “‘right’’ side) may be slightly larger than 
those on the other side of the rachis. However, if there is this ‘overdevelopment’ 
of the right side of a leaf, then the next higher leaf of the spiral often has its left 
side similarly overdeveloped, and the successive higher leaves reveal an alternation 
continuing this pattern. 


The nodes of the leaf rachis are closer nearer the leaf apex, and the internode 
distance is correlated approximately with leaflet width, being about the same or 
slightly more than the width of the nearest leaflets. The lower internodes elongate 
first and most rapidly. 


The stipules are short and connate into a narrow ligular rim, ciliate in the 
very youngest stage but soon glabrate. Internodes on the orthotropic shoots are 
so short that the sheathing bases of the petioles are in contact. 


The fruits are about 5-6 mm long, globose to broadly ellipsoid, capped by 
the remnant calyx rim and the short persistent conic style. 


Seedlings. Seedlings of this species are epigeal, with a pair of opposite bluntly 
ovate cotyledons about 1 cm long. The first several true leaves are simple, ovate- 
acuminate; these are eventually followed by pinnate leaves. 


Brassaiopsis Decne. & Planch. 
(Wardenia King, syn. nov.) 


The genus Wardenia with its single species W. simplex King has been found so 
far only in a few localities in Perak and Selangor and hitherto has been considered 
monotypic. F. R. I. Kepong Herbarium has several specimens of Wardenia simplex 
King, the type species of the genus cited below. These collections supplement the 
original from Ulu Kerling, Perak, made by Kunstler over 80 years ago. 


Two collections made in 1955 and 1969 in Trengganu, appear to represent a 
second species of this genus. It is described as a ‘fleshy treelet’ and differs from 
W. simplex in several obvious features, its elliptic acuminate cuneate (rather than 
ovate subcordate) leaves, much shorter and more slender petioles, and diminished 
inflorescence with little if any branching, and perhaps its longer pedicels and 
sparser puberulence. The material is rather incomplete; no fruits are available. The 
flowers are evidently past anthesis and the ovaries starting to enlarge. They clearly 
show a bilocular ovary, each cell with a single ovule. This conflicts with King’s 
Original generic description of Wardenia which calls for a unilocular ovary that 
develops a dissepiment later, separating the seeds, but agrees with the diagnosis 
of Brassaiopsis. According to Philipson (Bot. J. Linn, Soc, 63, Suppl. 1: 90, figs. 
1-6. 1970) the original observations may have been wrong as the type collection 
definitely has 2-locular young fruits. The floral character of “‘ovary 1-celled. 
2-ovulate”” should be treated with caution, however, as in any case it ‘disappears’ 
in the fruit phase which is more normal for Araliads, i.e. there is one seed per 
loculus. The ovular character may not differ therefore from the situation in 


Araliaceae 281 


Brassaiopsis Decne. & Planch. (including Fuaraliopsis Hutchinson. which is 
Araliopsis Kurz non Engler). It would appear that Wardenia differs from 
Brassaiopsis only in having simple leaves. However, simple-leaved species of 
Brassaiopsis have already been described. Philipson (1951)* for example describes 
B. castaneifolia from Burma, which has simple leaves. Thus the generic distinction 
does not appear to be tenable. The venation pattern in the simple-leaved species 
is quite easily reconciled with that of the compound or palmately lobed-leaved 
species. 

In both B. (Wardenia) simplex and B. minor, and in B. polyacantha and B. 
elegans, the young parts are covered with a rufous furfuraceous indument. 
Microscopic examination reveals that this is made up of numerous slender 
brachiate to stellate hairs with a variable number of branches and considerable 
variation in size. These structures are more or less ephemeral and the stems, 
leaves, etc., become glabrescent with age. These hairs are virtually identical in all 
four species, though their abundance varies. They are densest on the innovations 
and buds of B. polyacantha. 

The sheathing base of the petiole and the adnate stipules are similar in all 
four species. The petiole base is obliquely invaginated and the basal margins 
abruptly produced into a pair of lobes at a certain distance away from the base, 
as in shown in the accompanying figures. This pattern also shows up in the lowest 
inflorescence bracts, which appear to be much modified leaves with the blade and 
petiole reduced effectively to the sheath and ligule, although a prolongation 
representing the blade (and showing lateral nerves) but tightly cylindrical-subulate, 
may be present. 

The venation patterns in the leaves of all four species is comparable; the 
lowest lateral nerves are always inserted on the midrib by means of an outcurved 
junction. They are elevated beneath. There are comparatively few, i.e. widely 
spaced secondaries often with subparallel intermediate nerves between them. The 
lowest pair of lateral nerves tends to run up at or very near the blade margin and 
justifies the term ‘subtriplinerved’. In the palmate-leaved species these nerves give 
off leaflets which are thus asymmetrical. The overall veining pattern is campto- 
dromous. 

Consideration of these further characters, indument, petiole-ligule-stipules, and 
veining, adds weight to the decision to reduce Wardenia to Brassaiopsis. 

In R. Viguier’s anatomical study of Araliaceae (1906) Brassaiopsis was 
included (in the ‘Shefflerineés’ p. 96) and a characterization of its petiole anatomy 
was presented (the species not specified). For Wardenia, Viguier apparently had 
no material and he was unable to compare it anatomically with other genera. (He 
expressed doubt that it pertained to Araliaceae and suggested that it might belong 
to the Cornaceae). However, such doubt is unnecessary and it is preferable to 
reduce Wardenia to the status of a synonym of Brassaiopsis. This is also the view 
of D. G. Frodin (Misc. Rec. Fl. Mal. Found. III: 8, 1973). 


Key To MALAYAN SPECIES OF BRASSAIOPSIS 


1. Leaves palmately lobed or divided 
2. Leaves palmately 5—9-lobed, large to very large. to 60 cm wide; small 
EE ye tn SRE ec i eine fie aie ie ea ats B. polyacantha 
2. Leaves digitate with 3 or 5 or up to 3 leaflets, medium, to 25 cm. wide. 
3. Leaflets 3-5: inflorescence branches with terminal umbel and a 
ES EO OS Ae Or Ee ee oe B. elegans 


3. Leaflets 5—9: inflorescence branches racemose-umbellate ............ 
Se eeebirerrsis . givissd. . aatcawadadd. Wasik -% Reds Ous 3s. B. glomerulata 


* Phillipson, W. R. 1951. Contrib. to our knowl. of Old World Araliaceae. Bull. Brit. 
Mus. (N.H.) Bot. 1 (1): 1-20. 


282 Gardens’ Bulletin, Singapore — X XX (1977) 


1. Leaves simple. 
4. Leaves ovate-subcordate, to 35 cm long or more; petioles 


stout elongate, 15 cm long, 5 mm thick. ............ B. simplex 
4. Leaves elliptic, cuneate at base, smaller, to 25 cm long; petiole 
to 7 cm Jong. 2 mm COCK far, tt acotk ans cae B. minor 


ENUMERATION OF MALAYAN SPECIES OF BRASSAIOPSIS 


|. Brassaiopsis polyacantha (Wall.) Banerjee, Indian Forester 93: 341. 1967. 
B, palmata Kurz, J. As. Soc. Bengal 39, 2: 77. 1870. King, Mat. Fl. Mal. 
Pen. in J. As. Soc. Bengal 67: 61. 1898; Ridley, Fl. Mal. Pen. 1: 887. 
1922. 


Malayan collections: KEDAH: Gunong Inas F. R., Sg. Kepang, alt. 3000 ft., 
tree 40 ft. tall, 3 ft. girth, fruit faintly bluish with white spots, in terminal raceme 
2ft. long, 7.2.1968, Whitmore F RI-4639 (KEP); FRI-4644 (KEP). — Ulu Pantai 
Mulik, 22.3.1938, Sow KEP-34612 (KEP). PAHANG: Cameron Highlands, Gunong 
Brinchang, alt. 6000 ft., 4.5.1964, M.E.D. Poore KLU-8126 (KLU) — Telom Valley, 
21.8.1931, Jaamat KEP-25199 (KEP). — Bukit Tinggi village, Bentong Rd., tree 
30 ft., 8.2.1966, Kochummen KEP-97777 (KEP). — Bentong Rd. between Genting 
Sempah and Bukit Tinggi, alt. 1500 ft., 21.12.1968, Stone 8319 (KLU). — Bentong 
Rd. just beyond Genting Sempah, 1800 ft. alt., 2.2.1968, Whitmore FRI-4623 
(KEP). PERAK: Maxwell Hill Road, 54 miles, 29.10.1969, Kochummen F RI-2860 
(KEP), 

Poore 8126 differs in having more coriaceous leaves, appressed stellate hairs, 
thornier petioles. It may be a different species. 


2. Brassaiopsis glomerulata (BI.) Regel, Gartenfl. 12: 275. 1863. 
B. speciosa Decne, & Planch. Rev. Hort. 4, 3: 106. 1854. Hedera glome- 
rulata Blume, Bijdr. 1: 872. 1826. — Aralia glomerulata (Bl.) DC. Prodr. 
4: 265. 1830. 


Distribution: From India and S. W. China to Malaysia, Sumatra, and Java. 


MALAYA: PAHANG, J. H. Burkill & R. E. Holttum SF-7882 (SING); s.n. 
in 1922 (SING). NeGri SEMBILAN, Md, Nur SF-11729 (SING). 


This species may be recognized by its stature, for it becomes a tree up to 10 m 
tall; by its distinct (petiolulate) leaflets; and by the long inflorescence branches 
bearing umbellules of about 25-35 flowers. 


3. B. simplex (King) B. C. Stone, comb. noy. Wardenia simplex King, Mat. 
Fl. Mal. Pen. in J. As Soc. Bengal 67: 60. 1898. Ridley. Fl. Mal. Pen. 
1: 887. 1922. — Fig. 3. 

MALAYA: Perak, Slim Hills F. R., ridge top, 2500 ft. alt., unbranched 
small tree 15 ft. tall, trunk with distant thorns, inflorescence terminal, 4 Sept. 1966, 
Whitmore FRI-794 (KEP). SELANGOR, Compt. 12, Chadangan F. R., Ulu 
Selangor, hill forest, 1200 ft. alt., tree 10 ft. tall, girth 4 in., 27.9.1966, Chelliah 
KEP 98147 (KEP). 


4. B. elegans Rid]. Fl. Mal. Pen. 1: 888. 1922. — Fig. 4. 

MALAYA: PAHANG; Cameron Highlands, J. Wyatt-Smith s.n. (KEP). 
SELANGOR: Genting Sempah Ridge trail, 2500 ft. alt., 11.11.1970, Stone 9578 — 
(KLU). 


5. Brassaiopsis minor B. C. Stone, sp. nov. — Fig. 5. 


Arbuscula, ramulis 5 mm diametro. Folia simplices elliptica marginibus in 
partem mediam distalem serratis; petiolo 6-8 cm longo c, 1.5—-2 mm diametro basi 
breviter vaginantio; lamina tenue coriaceo basi subtriplinervio nerviis c. 6-8 


Araliaceae 283 


FRI-1274] 


Fig. 3. Brassaiopsis simplex (King) Stone. (Wardenia simplex King). Leaf, inflorescence, 


floral details, stipule detail, pubescence. FRI-12741. 


284 Gardens’ Bulletin, Singapore — XXX (1977) 


S 
= 
0 90 
4 9 
= D 
- *, 
A\ a ‘ s 
o hi Yee 
Om 
- A 
= 4 
YX . 
> 
« 
4 @ 
D 


Fig. 4. Brassaiopsis elegans Ridl. Leaf, inflorescence, floral details. Stone 9578. 


Araliaceae 285 


Fig. 5. Brassaiopsis minor Stone. Leaf, stipules, bracts (left: dorsal, right: ventral), 
inflorescence, and floral details (B':— the lowest inflorescence bract with a stipular- 


ligulate base). Type collection. 


286 


Gardens’ Bulletin, Singapore — XXX (1977) 


Fig. 6. Comparison of Schefflera singularis Stone with two Dendropanax species 
A:—Schefflera singularis, leaf and floral details, type collection. 


B:—Dendropanax maingayi King, leaf and floral details, KEP 66583. and fruit, 
Poore 1072. 


C:—Dendropanax lancifolius Ridl. cfr., leaf and floral details, Symington 25458. 


Araliaceae 287 


paribus, usque ad 25 x 12 cm, apicem versus abrupte acuminato, basi cuneato. 
Inflorescentia terminalis, brevis, 4-5 cm longis, bracteatis, pauci-vel non-ramosis, 
umbellatis; bracteolis deltoideis c. 1-1.5 mm longis rufo-puberulento-ciliolatis; 
pedicellis c. 15 mm longis; calyce 5-dentato; ovario rotundato-lobulato, depresso, 
biloculare, stylis binis unitis columna 1 mm longo formantibus. Loculi ovarii 2, 
uniovulati. 

Holotype: MALAY PENINSULA: TRENGGANU, Gunong Pahang, Ulu 
Brang, camp 3, humid deep valley on granite; alt. 3500 ft., fleshy treelet, flowers 
white; 21.9.1969, Whitmore FRI-12741 (KEP). 


Additional spec. examined: TRENGGANU: 34th mile. Kuala Trengganu, Besut 
Rd., lowland forest, (W. side), woody 2’ high, 8.9.1955, J. Sinclair & Kiah SF- 
40775 (SING). 


This is another simple-leaved species of Brassaiopsis. Originally I had con- 
sidered it as a new Wardenia. The flowers show clearly that the ovary is bilocular 
and there is a single ovule in each loculus. The slender shorter petioles, elliptic- 
cuneate leaves and shorter inflorescence quickly distinguish this from B. simplex. 


Hederopsis King 


Type species: Hederopsis maingayi King, Hk. FI. Brit. Ind. 2: 739. 1898. 


Originally described from rather inadequate material, this genus (endemic in 
Malaya and apparently Sumatra, monotypic) has in the last few years been 
collected rather widely and is in fact locally common in a few localities. It has been 
suggested by Frodin that the genus could be merged with Macropanax Miq. which 
appears to differ only in its usually 2-3-locular ovary, this structure in Hederopsis 
being normally 5-6-locular. The difference is a slight one, but King’s genus can be 
retained for the time being until a broader-based comparison can be made. The 
leaves are palmate-digitate, often with 5 leaflets, or with 3, or even 4, and sometimes 
(on flowering branches) reduced to 1 leaflet. The slender petioles may reach a 
length of 56 cm, and the terminal leaflet (always slightly longer than the laterals) 
a length of 31 cm and a width of 12 cm. The margins are dentate with small rather 
distant teeth. The petiolules may reach a length of 15 cm (again shorter on the 
lateral leaflets). — Fig. 7. 


The spirally disposed leaves have a rather short sheathing base about 1 cm 
long or less; when very young the margins are ciliate with pale brown multiseptate 
hairs, which are often laterally adnate in flat ‘bundles’. The juvenile stems are 
similarly rather densely hairy, but the hairs are ephemeral and all mature and 
submature shoots are glabrous. The base of the inner surface of the sheath shows 
a purplish spot adjacent to the axillary bud. The bud is compressed, purplish, 
pubescent, and bears 2 imbricate minute prophylls (which are more or less deltoid 
when flattened); these are equivalent to the leaf-sheath and show the ciliate margin 
and, often, a more or less produced apical elongation sometimes bearing three tiny 
lobes (suppressed leaflets). The outer face is usually rather pubescent. Similar 
prophylls occur on the terminal shoot in the uppermost region during renewed 
growth following the resting period, which is evident from the succession of several 
long internodes followed by a short series of very crowded internodes on the main 
branches and leafy shoots. 

Individual plants of this species become attractive small trees with a rounded 
crown, the trunk sometimes exceeding 90 cm girth, often fluted at base, giving 
off a few main branches; the whole tree may reach 15-20 m height. 

In Ulu Langat (Selangor) above the Reservoir (Ponsoon) there are several 
of these trees, copiously fruiting in January-February 1975, The fruits are sub- 
globose and still green when ripe. 


288 Gardens’ Bulletin, Singapore — XXX (1977) | 


Fig. 7. Hederopsis maingayi King. Leaf and floral details, and fruit. 


Araliaceae 289 


Spec. examined: MALAY PENINSULA: KeEpaH: Ulu Muda Forest Reserve, 
Compartments 115 & 116, 1500 ft. alt., a tree 40 ft, high, girth 24”, bole fluted, with 
small buttresses, bark rugose, lenticellate, inner bark thick brown, with watery 
exudate, wood white, fruits green. 21.1.1969, Y. C. Chan FRI-6782 (KEP). 
PERAK: Bintang Hijau F. R., secondary forest, tree 50 ft. tall, 26” girth, flowers 
pale yellow, faintly aromatic, cut bark smelling of resin, bole slightly fluted, 
22.11.1966, Rahim Ismail KEP-98531 (KEP). KELANTAN: Ulu Sungei Lebir 
Kecil, 500 ft. alt., 17.9.1967, P. F. Cockburn FRI-7123 and 7123 (KEP). SELANGOR: 
Ulu Langat, 2 miles from Kg. Injin Satu, tree 50 ft. high. 17.4.1963, Abdul Samat 
196 (KIU). — Ulu Langat Forest Reserve, tree 30 ft. high. 9.3.1966 Whitmore 
FRI-106 (KEP). — Ulu Langat, Bukit Lakwak, Kg. Panson, n.v. (Temuan) 
““Kayu pacha” 4.10.1958, Gadoh, Phytochem. Survey 1015 (KEP). — Ulu Langat 
above NEB works, stream valley, tree 10 m x 35 cm, fruits green, 7.2.1975, 
M. M. J. van Balgooy & B. C. Stone 2249 (KLU etc.). PAHANG: Ulu Tembeling, 
lowlands, 28.5.1931, Henderson SF. 24550 (SING). — Ulu Gali, Raub, 27.9.1929, 
Kalong KEP-20274 (KEP). — Cameron Highlands, K. Mensum to Boh Estate, 
24.9.1971, H. S. Loh FRI-17399 (KEP). — Same locale, 3.3.1972, alt. 2000 ft., 
tree 40 ft., girth 30”, Y. C. Chan FRI-19932 (KEP). 


Scheffiera J. R. & G. Forst. 
Schefflera singularis B. C. Stone, sp. nov. — Fig. 6A. 


Liana glabra innovationibus inflorescentiaque excepto, pilibus perminutis 
stellato-quinque-brachiatis; ramis usque ad 1 cm diametro teretibus cortice griseo; 
ramulis foliaceis usque ad 6 mm diametro in sicco longitudinaliter canaliculato- 
ruguloso; foliis simplicibus elliptico-obovatis marginibus valde sed anguste 
revolutis apice obtusis vel minute apiculatis vel rotundatis vel retusis, basi cuneatis, 
rigide percoriaceis costa in canaliculo costali utrinque leviter elevato, nerviis 
invisibilis; petiolo 5-20 mm longo. 1.5— mm crasso in sicco ruguloso basi ligulato 
ligulo 5 mm longo marginato; lamina 2-12 cm longo 1.5-6 cm lato apiculo 
0.5-2.5 mm longo vel nullo; nerviis lateralibus c. 5-7 paribus sed omnino obscuris, 
nerviis basalibus marginam formantibus. Inflorescentia terminalis bisumbellatis 
(composito-umbellatis) usque ad 3 cm longis axibus griseo-albide furfuraceis 
demum glabrescentibus bracteis deltoideis c. 4 mm longis bracteolis c. 1-2 mm 
longis scarioso-ciliato-marginatis. Umbellae c. 5-10-floriferae, pedicellis c. 
4 x 0.8 mm, calyce obconico 3 x 3 mm margine obscuriter 5-dentato minutissime 
ciliolulato, petalis 5 deltoideis 2 x 1.8 mm intus obscuriter carinatis apicibus 
unguiculatis, staminibus 5 filamentis albidis c. 1 mm longis antheris submedifixibus 
ovatis 1 mm longis erectis, disco leviter elevato, stylopodio late conico truncato 
vix 1 mm alto 2 mm lato, stigmatibus 5 obscuris radiatis ad apicem stylopodii 
sessilibus. Fructus immaturis glabratus obconico-turbinato-hemiglobosus 5 x 4 mm 
calyce undulato alte-marginatus stylopodio centrale 1 mm alto. 


Holotype: MALAYA: PaHANG, Gunung Ulu Kali, summit area, 5000-5800 
ft. alt., elfin forest with Dacrydium, big shrub with thick brittle leaves pale tawny- 
greenish beneath, 23.7.1967, B. C. Stone 7175 (KLU). 


Other specimens examined: MALAY PENINSULA, PAHANG, Gunung Ulu 
Kali, summit area, shrub in mossy forest, 19 Sept. 1967, J. Dransfield KLU-8068 
(KLU); same location, 27.6.1971, Mahmud b. Sidek and W. R. Stanton KLU- 
16312 (KLU). Ginting Highlands Rd. 4.5 miles (c. 3000’), 12.6.73, Mohd. Shah & 
Md. Ali MS. 2987 (climber? — fis. dark green in bud), (SING). Isotypes in 
I and KLU. 


290 Gardens’ Bulletin, Singapore — XXX (1977) 


This very distinct endemic species so far has been collected only on the 
summit area of the mountain Gunung Ulu Kali, in the Ginting Highlands of 
Pahang, on the Selangor border. At first it was considered to be a new species of 
Dendropanax but after much discussion and hesitation it is here described as a 
Schefflera. In nearly all features this species agrees with the formal definition 
of a Dendropanax, but differs in having a perceptible though limited ligular process 
at the base of the petiole, similar to that in at least most Scheffleras. In addition 
the exceedingly coriaceous leaf, with virtually invisible venation, appeared 
inconsistent with the condition in Dendropanax, which at least in the Malaysian 
species, is characterized by rather evident and subtrineryed leaves, as may be seen 
in D. maingayi and lancifolius, illustrated in Fig. 6 B and 6 C. Schefflera singularis, 
as shown in Fig. 6 A, not only has invisible nerves and evident stipular ligule, but 
a more oboval laminar shape than the two species of Dendropanax. Nonetheless, 
in floral characters there is almost no difference between the new Schefflera and 
the two Malayan Dendropanax species, as the figures cited also display. 


Within Schefflera the vast majority of species possess compound leaves with 
trifoliolate or palmate leaflet arrangements, Rarely, as in Schefflera lanceolata Ridl., 
simple leaves occur, though even in this species ojder branches may show 
trifoliolate leaves. Another such example is Schefflera simplicifolia Merr. of the 
Philippines. Among Malesian species I know of no other similar cases, though 
they may occur. Thus Schefflera singularis stands apart from nearly all other 
Malayan Scheffleras by its simple leaves. 


Another difference lies in the form of the inflorescence, which in Schefflera is 
usually a compound umbel, i.e. an umbel of umbels or umbellules, while in 
Dendropanax the umbel is often simple; but there are also species of Dendropanax 
with compound umbels. 


The genus Schefflera includes species which are climbers, others which are 
shrubs or trees, the latter often epiphytic. Dendropanax species are apparently 
mostly shrubs or trees. Schefflera singularis is a high-climbing woody liana. 


A comparison of generic diagnoses in such standard works as various floras, 
treatise* on the Chinese Araliaceae (Sargentia 2. 1942), and Vol. 2 of Hutchinson’s 
“Genera of Flowering Plants” (1967), does not appear to reveal any completely 
dependable differential features to distinguish Schefflera from Dendropanax except 
perhaps the presence of a developed stipular ligule. The degree of connation or 


separation of the style-tips seems to be highly variable within Schefflera and not 
a diagnostic character. 


In the light of these facts it is evident that further work is necessary to clarify 
the distinction between Dendropanax and the small number of Schefflera species 
with predominantly simple leaves. One may in fact question whether Dendropanax 
lancifolius Ridl. is not itself misplaced in Dendropanax. (In passing it may be noted 
that the generic name Dendropanax, although treated as neuter by the original 
authors, has usually been considered masculine in gender, and thus the epithet is 
spelled here Jancifolius rather than lancifolium as it appears in Ridley’s Flora). 


Acknowledgements 


_it is a pleasure to record the debt owed to Prof. W. R. Philipson of the 
University of Canterbury for stimulating discussions concerning Araliaceae and 
for his several suggestions which have improved and augmented this paper. I am 
also grateful to D. E. Frodin, University of Papua New Guinea, and Prof. 


? Bye. i. Li. 


Araliaceae 291 


I. V. Grushvitzky, Komarov Botanical Institute, Leningrad, for their helpful 
advice. Thanks are also due to the Director and Curators of the Botanic Gardens, 
Singapore, and the Forest Research Institute, Kepong, Malaysia, for the opportuni- 
ties to study the specimens of Araliaceae in their care. Shamsuddin bin Abdul 
Jalil, a former Botany Honours student in the University of Malaya, assisted in 
this work by carrying out a useful preliminary study of the Malayan species of 
Arthrophyllum as an Honours thesis under the writer’s supervision, and his 
contribution is gratefully acknowledged. 


Reference 


Philipson, W. R. 1970. The Malesian species of Gastonia. Blumea 18: 490-495. 


nd cane r 
ler aoigeqe: sry ot Ls Nee 


vik hee apie Tequy,..4 Tail, 


aie 
‘ edecr ey. 


Tere UL 


we 
Case ey 
eR pani “ 


ehh t 
ibs 


he — ion 


ax. 


othe ety a: 


A Note on the Cytology of Botrychium lanuginosum 
and the Occurrence of the Genus in Malesia 


by 


A. C. JeERMy* and T. G. WALKER** 


Summary 


Over the past twenty-five years intensive plant collecting has shown two species of 
Botrychium (B. lanuginosum Wall. ex Hook. & Grev. and B. australe R. Br.) to be present in 
eastern New Guinea, the former hitherto confined to Central—S.E. Asia and reaching to the 
Central Himalaya and the latter Australasian with a possible representative in S. America. A 
further species, B. daucifolium Wall. ex Hook. & Grev., is distributed from Sri Lanka 
(Ceylon) to the Philippines and Sulawesi (Celebes) with questionable outlying records in Fiji 
and Samoa, reaching S. China in the North, and replaced by a taxon of doubtful specific 
identity in Japan (B. japonicum (Prantl) Underw.). B. lanuginosum was found in large quanti- 
ties in the kunai grasslands at 1600 m in the Finisterre Mountains of E. New Guinea and 
material fixed in the field has proved to be octoploid m = 180) and also hexaploid, 
suggesting the tetraploid cytotype recorded by other workers in India and Sri Lanka may 
also be present in New Guinea. 


INTRODUCTION 


The appearance of Botrychium in S.E. Asia and Malesia is seasonal. Its 
ecological requirements are little known but its appearance may be linked with 
the increased rainfall seen just before the full monsoon develops. For this reason 
limited collections have been made as the aerial parts of the leaf quickly become 
yellow and die down after the spores are shed. Three species have so far been 
recorded in Malesia, B. lanuginosum Wall. ex Hook. & Grev., B. daucifolium Wall. 
ex Hook. & Grev. and B. australe R. Br. The first and last species are components 
of well drained soil in the open anthropogenic grasslands in Malesia (from 
1600 — 2700 m in New Guinea) and on similar grassy banks or more rarely in 
woods on the Asiatic mainland. B. daucifolium is a species of more shady situation 
often in open woodland in the Himalaya and on the open floor of the rain forest 
in Borneo and Sulawesi. Betty Molesworth Allen (1959) discovered a colony at 
1500 m on Gunong Perdah in an opening in the tall forest, growing in a scrub of 
ogg i and young Cyathea. She records the soil as being a black sticky 

umus. 


KEY TO MALESIAN SPECIES OF Botrychium 


1 Fertile segment arising from the rachis of the sterile blade, ++ equal in length 
to.adjament. sterile pinnac ........ 000? aaah onan atee....... 1 lanuginosum 


1 Fertile segment arising from the petiole below the sterile blade; the stalk of 
the fertile segment 1 — 2 times as long as the sterile blade 


2 Fertile stalk + equalling sterile blade; upper pinnae linear-lanceolate or 


triangular, pinnatifid or dentate only ....................0.ccccceee eee 2 daucifolium 
2 Fertile stalk at least twice as long as the sterile blade; upper pinnules ovate, 


Se kv eer ne EA Pep, ee \ nee 3 australe 


* Botany Department, British Museum (Natural History), London, England. 
** Department of Plant Biology, University of Newcastle upon Tyne, England. 


293 


294 Gardens’ Bulletin, Singapore — XXX (1977) 


Cytology, Botrychium 295 


1 B. lanuginosum Wall. ex Hook. & Grev. Icon. Fil. tab. 79 (1829). 


The name first appeared in Wallich’s Catalogue of 1828 without a description 
and was described by Hooker, the figure by Greville (/.c.) drawn from material 
supplied by Wallich. Ballard (1940) discusses the typification and describes the 
specimen in Greville’s herbarium at the Royal Botanic Garden Edinburgh from 
which the two lower pinnae were removed, thus giving rise to the abnormal! plant 
seen in his plate 79. 


The species ranges from the Central Himalaya across China to Taiwan, south 
to Sri Lanka on the Indian subcontinent and in Malesia to Luzon, Sumatra, Java 
and New Guinea. The species is distinct in having a broadly deltate tripartite 
sterile blade, the lobes usually 3-4 pinnatisect, and with the fertile scape 
arising from the rachis above the insertion of the two lower pinnae (see Figs 
1 & 2). The bud, rachis and especially the costal axils are usually sparse to 
densely covered with long unbranched hairs although some fronds in the New 
Guinea population mentioned below were almost glabrous. Hayata (1914) has 
described the glabrous form as B. leptostachyum, later to be combined by Nakai 
(1925) as a variety of B. Januginosum, but in view of the wide variation in the 
above population both are of doubtful status. The degree of dissection of the sterile 
blade varies, younger plants being usually less compound, but always more 
dissected than either B. daucifolium or B. australe. There is no apparent correlation 
between ploidy level and degree of cutting or size of sterile frond (see Figs 1 & 2). 


2 B. daucifolium Wallich ex Hook. & Grev. Icon. Fil, 2, tab. 161 (1830). 


Wallich (1828) listed the first gatherings of this (No. 49) as B. subcarnosum 
and later by the name Hooker took up for it. The specimen drawn is in the 
Edinburgh herbarium and other plants of the same gathering at the British 
Museum (Natural History) London, 


The species ranges from a similar area in the centra) Himalaya east to Taiwan 
and south to Sri Lanka. In Malesia it is found throughout the archipelago to 
Sumbawa Is. (G. Batulanteh), Celebes (G. Bonthain) and Mindanao (Mnt Apo) 
and Negros (Cuernos de Negros). Allen (1959) found two populations in the 
Cameron Highlands area of Pahang. There are specimens in the Kew Herbarium 
from Upola, Samoa and Nadavivatu, Fiji which compare well with material of 
B. daucifolium from Asia, but a closer study of more recent material needs to be 
made. 


B. daucifolium is distinct in having thin, but not membranous, lamina tissue 
and a less dissected frond in which the upper pinnae are barely cut (see Fig 3) 
but lobed with the segments toothed. The fertile spike arises from the petiole about 
10 cm or more below the insertion of the blade and the fertile segments are strict, 
not spreading as in the other two species. The species varies in its hairness, some 
species in Borneo being quite densely pubescent on the rachis and costae, Allen 
(1959) mentions that the very young fronds on her Malayan plants were covered 
with copious pale hairs. Himalayan plants tend, in herbarium specimens at least, 
to be more glabrous, as do the adult plants of Allen, who gives a fine general 
photograph of the species (/. c. facing p. 251). 

Material from Japan, E. China and some from Taiwan, being more mem- 
branous, more deeply toothed and with the fertile stalk somewhat longer, has been 
separated off as B. daucifolium var. japonicum Prantl, raised by Underwood 


Facing page 
Figs. 1-4. Silhouettes of Botrychium. 1 & 2. Silhouettes of Botrychium lanuginosum from 
Moro, Finisterre Range, New Guinea showing range of dissection. 1: Jermy 3887 


(x 4); 2: Walker 8333, octoploid plant from same population (x 4). fs = fertile spike. 
3: B. daucifolium Gunong Dempo, Sumatra, C. J. Brooks 15881 (x 4). 4: B. australe 
Western Highlands District, New Guinea J. S. Womersley, NGF 15213 (x 4). 


296 Gardens’ Bulletin, Singapore — XXX (1977) 


(1898) to a species. It is certainly related in its morphology to the typical mainland 
asian B. daucifolium. A small form was described by van Alderwerelt van 
Rosenburg (1911) from Preanger Regencies, Java under the name of var. parvum. 
Later in 1913 the same author described yet another variety, subbasalis, which was 
not only smaller than usual for B. daucifolium but had the fertile spike arising 
towards the base of the stipe as in B. ternatum and B. australe. Clausen (1938) 
considered this, from the published description only, to be a possible new species. 
We have not studied van Alderwerelt van Rosenburgh’s type and must reserve 
judgement but it is our experience that juvenile and depauperate Botrychiums can 
appear very abnormal. 


3. B. australe R. Br. Prod. Fl, Nov. Holl.: 164 (1810). 


Collected and described by Robert Brown from material from Tasmania and 
New South Wales. The specimen in the herbarium of the British Museum (Natural 
History) from Parametta, Port Jackson can be taken as typical but the specimen 
at Kew is more complete (it contains roots) and is best to be designated lectotype. 
The species ranges from South Austra'ia to New Zealand and north into south 
Queensland where it was recorded.as B. ternatum (Thunb.) Sw. by Bailey (1883). 
It differs from the latter in its dissection of the thicker lamina which has hyaline 
marginal cells. 


The sterile leaf is 10-20 cm long, ternate, with the central leaflet larger and 
more compound. The upper pinnae of this leaflet are ovate and cut to the costa, 
the resulting segments glabrous, linear and dentate at the truncate apex; the tissue 
is fleshy and the lamina margin is made up of two to three rows of hyaline cells. The 
compoundly branched and spreading fertile segment arises in the lower half of the 
stipe to a height of 30 cm. (see Fig. 4). 

The map in Fig. 5 shows the range of the species in Australia. Its presence 
in Papua New Guinea in the Southern and Western Highlands Districts is a logical 
extension of this range. It grows in high altitude anthropogenic grasslands 
(Miscanthus-T hemeda associations) and around 2700 m in Danthonia-Poa grass- 
land and Cyathea scrub and in two localities together with B. lanuginosum. The 
taxonomy of this australasian complex is little understood and is being reviewed 
by New Zealand workers at present and their results are awaited with interest.* 


THE CyToLocy oF B. lanuginosum IN NEw GUINEA 


Whilst collecting fern material for subsequent taxonomic and cytological studies 
in Madang District, east New Guinea in 1964 the authors showed children in 
Moro, a village at 1680 m at the head of the Gusap River in the Finisterre 
Mountains, an illustration of Botrychium, Within the hour the children returned 
with armfuls of B. lanuginosum collected from amongst the kunai (Imperata 
grass) on old garden sites. This is a good example of the dangers of this method 
of collecting easily recognisable but possibly rare species. Luckily this population 
could withstand such collecting and some 50 specimens were taken to show the 
range of variation. Seven specimens thus collected had meiotic material and were 
fixed in the field. 


One specimen (Walker 8333) showed exactly 180 bivalents at meiosis (Plate 1) 
and three others (Walker 8332, 8334, 8336) 270 chromosomes and hybrid meiosis. 
The base number for Botrychium is well established as 45 and thus these New 
Guinea specimens were octoploid and hexaploid respectively. This contrasts with 
other counts for the species from the Indian subcontinent where Verma & 
Loyal (1960) recorded n=90 in a Himalayan specimen and Ninan (1956) 


a Se Re RENN Mintle a 
* Dr J. Braggins, University of Aukland, N.Z., is of the opinion (pers, comm., 1976) that the 
New Guinea specimens constitute a new subspecies. 


Cytology, Botrychium 297 


recorded n=90 and 2n=180, also for N India; Manton and Sledge (1954) showed 
a similar situation in Sri Lanka (n=90). Hence all previous counts are at the 
tetraploid level. The fact that we have hexaploid material showing hybrid 
behaviour suggests that the tetraploid may also be present in New Guinea. 


) 


3 


ee ee ee Se ee ee ee 
. 


B. lanuginosum 
B. australe 


5. Distribution of the three species of Botrychium found in Malesia. 


298 Gardens’ Bulletin, Singapore — XXX (1977) 


REFERENCES 


Allen, B. Molesworth (1959) Malayan Fern Notes, A preliminary note. Gard. 
Bull. Singapore 17: 251-252. 

Bailey, F. M. (1883) A synopsis of the Queensland Flora. Brisbane. 

Ballard, F. (1940) Notes of ferns and fern allies IJ. Kew Bull. 1940: 29-31. 


Clausen, R. T. (1938) Monograph of the Ophioglossaceae. Mem. Torr. Bot. Club 
19 (2): 3-177. 


Hayata, B. (1914) Icones Plantae Formosae 4: 134. Taipei. 


Manton, I & Sledge, W. A. (1954) Observations on the cytology and taxonomy 
of the pteridophyte flora of Ceylon. Phil. Trans. roy. Soc. Lond. 283B: 127-185. 


Ninan, C. A. (1956) Cytology of Ophioglossaceae. Curr. Sci. 25: 161-162. 
Nakai, T. (1925) Bot. Mag. Tokyo 39: 192. 


Underwood, L. W. (1898) American ferns I. The ternate species of Botrychium. 
Bull. Torr. Bot. Club 25: 521-541. 


Van Alderwerelt van Rosenburg C. R. W. K. (1911) New or interesting Malay 
ferns 3. Bull. Jard. Bot. Buitenz. ser. 2 (1): 3. 


(1913) New or interesting Malay ferns 5. ibid. ser 2 (11): 6 et 
tab. 3. 


Verma, S. C. & Loyal, D. S. (1960) Chromosome counts in some ferns from Naini 
Tal. Curr. Sci. 29: 69-70. 


Wallich, N. (1828) A numerical list of dried specimens of plants in the East India 
Company museum collected under the superintendence of Dr Wallich, London. 


; a 0 ge iso? . 
el eK. 
cules 8 ete 4 
4 eats 
4, *§ 5 : - 
ey » ‘4 
v i 1 ee % ieee 
4 ‘ : . 
ee wee 
s. a poet. x @ sole 
> if 
a 
yr uta® a a 
- hee, Adee 
es 4 ad 
2 #3 > . ‘ 
se 8 |. © 4 " 
® ’ oe 
= xX 
on %, e_. 
Sete ® § 
"4 thks 
e e 
é : te 


Te oe 
_& 

: 
“ts 


Plate 1. Permanent aceto-carmine squash of metaphase I of a spore mother cell of Botrychium 
Ranugeniosts (Walker 8333) together with explanatory diagram below. Magnification 
1000. 


A Revision of the Malesian Species of Arthrophyllum BI. 
(Araliaceae) 


by 


W. R. PHILIPSON 


Department of Botany, 
University of Canterbury, 
New Zealand. 


The genus Arthrophyllum forms a well-defined group of species within the 
Araliaceae. Its principal distinctive characters are the single-celled ovary and the 
arrangement of the umbels on specialized flowering branches. The vegetative 
shoots bear pinnate or bipinnate leaves in spiral phyllotaxis. Branches do not 
develop until flowering occurs, when a terminal] tuft of lateral branches develops. 
These bear opposite leaves of reduced size and terminate in a whorl of secondary 
branches which may themselves bear umbellules or which may branch again 
before doing so (fig. 1). Vegetative growth is continued by one or more innova- 
tions, with spirally arranged leaves, arising below the crown of flowering branches. 


The geographical range of the genus almost coincides with that of Flora 
Malesiana, but extends slightly further west to the Nicobar Islands and northwards 
into Laos and Thailand, and the range does not include the Lesser Sunda Islands. 
Apart from the treatment in Ridley’s Flora of the Malay Peninsula, no revision 
of the species has appeared. The many taxa given specific names by early investi- 
gators (most of which are here regarded as synonyms) were never critically 
compared, so that knowledge of the genus has remained confused and fragmentary 
and botanists have tended to avoid it. Consequently it has been found necessary 
to describe several new species, while at the same time reducing several others 
to synonymy. 

The present revision is regarded as only tentative. Herbarium material is 
abundant, but most specimens are fragmentary and therefore fail to illustrate the 
_unusual branch-systems which characterize some species. Specific distinctions 
seem to be unusually difficult to define on the basis of herbarium specimens, so 
that knowledge of the group will remain superficial until a comprehensive field 
study can be undertaken throughout the range of the genus. 


The genus comprises two species-complexes, and also the anomalous 
A. proliferum. One of these complexes includes seven shrubby species with simpler 
inflorescences (no more than three degrees of branching), namely, A. maingayi, 
A. montanum, and A. alternifolium (from the Malay Peninsula, with the first 
species also from Sumatra and Borneo); A. papyraceum (from Sumatra); A. 
pulgarense and A. cenabrei (from the Philippines); and A. kjellbergii (from 
Celebes). The second complex consists of nine species of tree stature with larger 
inflorescences (having four degrees of branching). This complex occurs commonly 
throughout the range of the genus, and its subdivision into species presents con- 
siderable difficulty. To the east A. macranthum is abundant on the mainland of 
New Guinea, while on islands to the north a second species, A. pacificum, is 
recognized. In the Philippines and northern Moluccas this complex is represented 
by A. ahernianum, a species which also extends into the extreme north-east of 
Borneo. The complex is more difficult to classify further west and south, that is, 
in Celebes, Borneo, Java, Sumatra, and the Malay Peninsula. In the present 


299 


Gardens’ Bulletin, Singapore — XXX (1977) 


300 


‘(OI/1 x) 8uryouesg jo ‘(O€/1 X) seyouesq SULIIMOY Pue datezaSoA 
saoIdap Inoj YIM ‘jooys ZulIOMOY 9[3uIs suIMOYS JIQeY jo Yo}0¥s NjVWWILISVIP.. “wn70fissaaip unyxydosyjsp *] “B1y 
B JO SUIMLIP SIVeUIDYOS ‘wnNOfissaaip “py ~ ‘S14 . : 


DWYONN i) i> 
“22a | LESS L 
PAKS | 
DEG 9, 
ris we 
SEZ SU ZINE ome a 
Ly “Ahn Ee” 
\ Y [) 4 SQ YX = WFO ps 
wm | ~S SQ 1 AMA 
4S ; Ye yw 
cops RS LAS 
3 ST PVM 
= Ce AY NY 
y, HY OY S&S 
SK SS OY Y 
Z\ SF PS AY 
re eh) [Y SL . ; ey SY 2 
. cae Q Se Yi t ey : 


Arthrophyllum ‘301 


revision, in addition to the widespread A. diversifolium, five entities from this 
area are given specific rank: three species endemic to Borneo (A. crassum, A. 
collinum, and A. ashtoni; one most distinctive species, A. angustifolium, which 
is rare in Borneo and the Malay Peninsula; and a species, A. engganoense, which 
is known only from the small island, Enggano, off the west coast of Sumatra. The 
remaining great mass of material is here treated as a single species, A. diversifolium. 
The variation and nomenclature of this widespread taxon is discussed separately 
under the section on that species. 


ARTHROPHYLLUM 


Arthrophyllum Bl. Bijdr. Fl. Ned. Ind. ( 1826) 878. Mormoraphis Jack ex Wall. 
Cat. n. 4931 (1831). 


Sparingly branched trees or shrubs, unarmed. The leaves on vegetative shoots 
spirally arranged, imparipinnate, those on flowering branches often opposite, 
smaller, or reduced to a single leaflet; stipular sheath small. Inflorescence con- 
sisting of compound umbels, either solitary and terminal or more commonly borne 
on a cluster of specalized leafy branches arising from the axils of the uppermost 
leaves; pedicels not articulated. Flowers hermaphrodite; calyx an undulate rim; 
petals 4-6, valvate in bud; stamens 4-6, anthers curved, basifixed; ovary turbinate, 
l-celled; disk fleshy, rising in the centre to the + sessile capitate stigma. Fruit 
ovoid or spheroidal, often oblique; exocarp leathery; endocarp cartilaginous; seed 
solitary, pendulous; endosperm deeply transversely ruminate. 


About 18 species (17 in Malesia) extending from the Nicobar Islands and 
in the larger islands or regions is as follows: Sumatra 4; Malay Peninsula 5; Java 
1; Borneo 7; the Philippines 3; Celebes 2; Moluccas 2; New Guinea and Bismarck 
Archipelago 3. The genus Eremopanax, with several species in New Caledonia, 
may be congeneric, 


KEY TO THE SPECIES ... 


1. a. Inflorescence becoming paniculate by the successive development of 


branches below the umbellules (figs. 3, 4) .................. I. A. proliferum 
ge Be Bl) Ren ee rn oe 2 
2. a. Inflorescence with four orders of branching (fig. 2) as ahes DE No 3 


b. Inflorescence with three (or fewer) degrees of branching (figs. 5, 6) 


PCRS SETH SEES EEE HEHH SHS EHEHEHHE HEHEHE EHH HET HEHTHEHEHEHESHE HEHEHE HEHEHE HEHEHE HEHE HEHEHE HEHEHE HHH HEHEHE HEB E ES 


ee ec dha cr asncescccserccnnacusvscceseccee ce 4 
b. Leaflets broader rie Re habia oe nares Sct diiet ries , ‘delieenednante 5 
4. a. Leflets c. 4-7 cm NN OR ok ai ek. Se CO TIC 2. A. ashtoni 
b. Leaflets much longer ............0....66, NRIs. 3. A. angustifolium 
5. a. Pedicels (at early anthesis) + 10 mm long .................ecceeeeeee ees 6 
b. Pedicels (at early anthesis) c. 5 mm long, or shorter Maye Be, 7 


6. a. Peduncles of umbellules with bracts or bract-scars (fig. 13) .........:00008 
AO Elie nck eels ate ANE Vilas care Fp eyigs pF 8S Cunt beg Hew ps Shaye gd since y «fp 4. A. ahernianum 


b. Peduncles of umbellules (fig 14) without bracthint ‘goassieivol.wil........ 
Sich bedns 6s xs aeteRNGEe SMO) die Ws. UL. JAI. BALI 28. ‘5. A. engganoense 


302 Gardens’ Bulletin, Singapore — XXX (1977) 


7. a. Leaves associated with the umbels rotund (fig. 17) ..............:esueseeeeees 
yp... o). .nenio8. ol aunehor. zvicage. oauds..:Ated. aes 6. A. collinum 

b. Leaves (or leaflets) associated with the umbels ovate or elliptic ......... 8 

8. a. Leaves associated with the umbels ovate, + fleshy, with the lower surface 
smooth (lateral veins obscure) (fig. 18) .................. sence 7. A. crassum 

b. Leaves (or leaflets) associated with the umbels + elliptic, coraceous or 
chartaceous, veins ViISIDIC © <24........ :catenea..ceesssasegeuaceesees cane 9 

9. a. Young parts glabrous ....../....0i tide cesee eee ece scene 8. A. pacificum 
b. Young parts with rufous tOmentuM .........05...ceseee separ -cosceenezecrsens 10 


10. a. Umbels at anthesis with numerous filamentous pedicels (fig. 15) ......... 
iniawas. ao. 22ve0!. oAT:: bomany-.acdwade 26-4808 ben 9. A. diversifolium 


b. Umbels at anthesis with fewer stout pedicels (fig. 16) ...................0.05. 
FT SSI? | BOS , SOLOS tee her ton Soe eect 10, A. macranthum 


11. a. Leaflets membranaceous or chartaceous ................cceeee eee eee ene en ees » he 
b: Leaflets coriaceous 2) 0/ 00225. 2002.2... PRP. 4001, GE ae ... 14 

12. a. Mid-leflets about 16 cm long ................c cece seen ees 11. A. papyraceum 
b. Mid-leaflets + 8 cm long or shorter ............... cece cece ee eeeeeeeeneenes 13 

13. a. Leaflets usually 5-7 (fig. 7), primary inflorescence branches usually 
short (+ 3-6 cm) and without articulations ............... 12. A. maingayi 

b. Leaflets more numerous (fig. 10), primary inflorescence branches longer 
(10-20 cm) with one or more leafy nodes ............... 13, A. kjellbergii 

14. a, Leaflets c 5: ...celi iio eee 14. A. cenabrei 
b. Leafléte more nWNCTOUS 55 Feats oan senda de atesagtds o-oo vnunet peeaiee sail ae 15 

15. a. Petals & stamens B>.. .<cpts <i;44-. sis erseshilionan 15. A. pulgarense 
b.... Petals & stamens. 4. Of) Bic, 103s coadesbomsvesss she ¥t “es rdead nadienea eee 16 

16. a. Leaflets 6 cm long, or longer (fig. 8) .................000. 16. A. montanum 
Leaflets shaster CBR) 39) sib dantbchocnpatiebhisebonds 17. A. alternifolium 


1. Arthrophyllum proliferum Philipson, sp. nov. 


Arbor foliis imparipinnatis. Inflorescentiae ex suis propriis ramis natae, 
paniculatae factae ramis sub umbellulis iterum atque saepius ramificantibus. | 


_ Medium sized tree, glabrous, Leaves of the vegetative stems spirally arranged, 
imparipinnate, multijugate, 90 x 24 cm, of the flowering branches smaller with 
fewer pinnae or usually simple; leaflets obovate-oblong, c. 16 x 7 cm, chartaceous, 
margin entire, slightly revolute, base broadly cuneate to truncate, sometimes 
oblique, apex acute, Inflorescences terminating specialized plagiotropic shoots, 
bearing axillary flowering branches and ending in umbellules of a few flowers 
below which pairs of whorls of branches conitnue the growth of the inflorescence 
to produce an elongated panicle of umbellules. 


_ Endemic to Papua New Guinea. Type: Territory of New Guinea, Morobe 
District, Mauwata Banda L.A., Bulolo, Havel and Kairo NGF 17172 (Canberra). 


In mid-mountain rain forest, reaching the canopy, on steep slopes, 300-1200 m. 


The flower and fruit are typical of this well-defined genus, but the branching 
of the inflorescence is unlike that found in all other species. 


Arthrophyllum 303 


Figs. 3-6. 3. A. proliferum; a portion of inflorescence in the fruiting state (x 4); 
4, same, showing detail of ultimate branches of the inflorescence in the flowering 
stage (x 1). 5. A. maingayi, terminal inflorescence (x 4). 6. A. montanum, 
a single flowering shoot (x 4). 


2. Arthrophyllum ashtoni Philipson, sp. nov. 


Abor gracilis tomento fusco. Folia imparipinnata, foliolis lanceolatis vel 
late-lanceolatis, 3-7.5 x 1-2 cm. Inflorescentia ramis in proporium officium 
mutatis et in coronam terminalem dispositis, quarter ramificantibus. 


A slender small tree, to 5 m high, young parts with brown scurfy tomentum. 
Leaves of the vegetative stems spirally arranged, imparipinnate, multijugate, to 
about 30 cm long; of the flower-bearing branches smaller with fewer pinnae, or 
simple; leaflets lanceolate to broadly-lanceolate; 3-7.5 x 1-2 cm, thinly coria- 
ceous, margin entire, revolute, base broadly cuneate, apex tapered to sub-caudate, 
veins channelled above, visible below. Inflorescence a terminal cluster of specia- 
lized leafy branches; the main rays variable in length in the same inflorescence, 
ending in a whorl of secondary rays subtended by simple leaves; the secondary 
rays 4-12 cm long, bearing simple leaves, usually in an opposite pair and 
terminating in compound umbels; umbellules with about 8-10 flowers pedicels 
4-10 mm long, furfuraceous, 


Endemic to Borneo. Type: Brunei: Pagon ridge, Ashton Brun 2341 (Kew). 


304 Gardens’ Bulletin, Singapore — XXX (1977) 


Moss forest on sandstone ridge, and Kerangas forest, 1000-1550 m. 


The small narrow leaflets are very distinctive. No other species with small 
leaflets has inflorescences which branch to the fourth degree. 


3. Arthrophyllum angustifolium Ridley in Journ. Fed. Mal. St. Mus. 10 (1920) 


136. 


A shrub or small tree, 5 m high, young parts rufous tomentose, becoming 
glabrous. Lower leaves spirally arranged, imparipinnate, multijugate, about 
70-90 x 30-40 cm; upper leaves reduced mostly unifoliate, opposite, broader, 
with petioles about 2-4 cm long; leaflets coriaceous lanceolate, c. 15-22 xX 
1.5-2.5 cm, tapering to an acute or obtuse apex, base cuneate, margin entire, 
slightly revolute. Inflorescence a terminal cluster of specialized leafy branches, 
the main rays 30 cm (or more) long, bearing simple leaves in opposite pairs 
ending in a whorls of c. 10-12 secondary rays subtended by simple leaves; the 
secondary rays 8-12 cm long, terminating in an umbel of 5-12 tertiary rays c. 
2-3 cm long, each bearing an umbellule of c. 8-12 flowers, pedicels c. 5 mm long: 


Malay Peninsula and Borneo. Type: Perak, Gunong Keledang, Ridley 9683 
(SING). 


Occurs in forest and old regenerated forest on peat swamp at low altitudes 
or on ridges. 


The lanceolate leaflets are unlike those of any other species, The grey bark is 
minutely fissured and bears many small orange lenticels, The wood is soft and 
white. The Peninsular and Bornean specimens are similar, except that the flower 
buds are larger in the Brunei plant. 


4. Arthrophyllum ahernianum Merr. in Philipp. Journ. Sci. 1 (1906) Suppl. 109. 


— ‘tA. pinnatum Clarke” F. -Vill. Novis. App. (1880) 103 — Macropanax sp. 
Vidal Rev. Pl. Vasc. Philip. (1886) 145. — A. sablanense Elmer in Leafl, Philipp. 
Bot. 1 (1908) 331. — A. borneense Merr. in Univ. Calif. Pubs. Bot. 15 (1929) 
231, non Baker. — A. elmeri Merr. in Webbia 7 (1950) 319. — A. merrilliana 
Furtado, in Gard. Bull, 19 (1962) 185. 


Tree up to 15 m high, young parts with rufous tomentum., Leaves clustered 
at the ends of the branches, spirally arranged on the vegetative shoots, impari- 
pinnate, multijugate, articulated at the insertion of the pinnae; up to 200 x 60 cm; 
of the flower-bearing branches smaller with fewer pinnae, or simple; leaflets 
ovate-oblong, occasionally oblong-lanceolate, up to 35 x 12 cm, membranaceous 
or chartaceous, margin entire, revolute, base cuneate to rounded, usually oblique, 
apex short accuminate, Inflorescence a whorl of specialized leafy branches forming 
a terminal crown; the main rays up to 150 cm long (or more), bearing pinnate 
leaves usually in 1-2 opposite pairs, ending in a whorl of secondary rays up to 
30 cm long terminating in compound umbellules; umbellules with about 10-20 
flowers; pedicels c. 10 mm long (at anthesis). | | 


Extending from northern Borneo, throughout the Philippines to the northern 
Moluccas. Type: Lamao Forest Reserve, Luzon, Meyer 2780. 


Primary and second-growth forest, from lowlands to 1000 m. 


This .species replaces the more westerly A. diversifolium which it closely 
resembles. It is characteristically larger in all its parts, particularly in the size of 
the individual flowers and the length of their pedicels. There are fewer flowers in 
an umbellule. The distinction between these two species is not always easy to 
make, especially when the material is fragmentary. A few specimens from the 
Philippines e.g. Clemens 585, Merrill 1232 appear very similar to A. diversifolium, 
and it is possible that this species extends beyond Borneo, However, at present 


Arthrophyllum ri 305 


these specimens are regarded as part of the range of variation of A. ahernianum. 
Similarly, at least one specimen from southern Borneo (Buwalda 7973) approaches 
A. ahernianum in appearance. 


5. Arthophyllum engganoense Philipson, sp. nov. 


Arbor foliis imparipinnatis. Inflorescentia ramis in proprium officium mutatis 
et in coronam terminalem dispositis, quater ramificantibus; umbellulae pedicellis 
circa 5-10, c. 10-15 mm longis sub anthesi. 


A tree to 21 m high, becoming glabrous. Lower leaves imparipinnate, 
multijugate, 60 cm long or more, petiole 24 cm long, 6 mm wide, petiolules 
10-18 mm long; leaflets broadly elliptic to elliptic-oblong, c. 12-15 xX 6~7 cm, 
base rounded with a short asymmetrical cuneate centre, apex shortly apiculate, 
margin entire often undulate chartaceous. Flowering branches c. 40 cm long; 
leaves opposite, simple or unifoliolate, ending in a whorl of. simple leaves 
surrounding a compound umbel to 30 cm diam.; secondary rays about 8, c. 
10-15 cm long, tertiary rays c. 8, slender c. 20-40 mm long, without bracts; 
pedicels c. 5-10 per umbellule, c. 10-15 mm at anthesis. 


South Sumatra, Enggano Isd. Type: Liitjeharms 4260 (Bogor). 
Occurs in forest at low altitudes (up to about 100 m). 


The two known collections of this species are very similar and contrast with 
the widespread A. diversifolium because of the few flowered umbelles with long, 
spreading pedicels. 


6. Arthrophyllum collinum Philipson, sp. nov. 


Arbor parva in prima aetate tomento rufo. Folia imparipinnate, foliolis 
oblongis, late ellipticis vel rotundis. Inflorescentia ramis in proporium officium 
mutatis et in coronam termnalem dispositis, .quarter ramificantibus; bracteis 
rotundis, coriaceis. 


Sparingly branched shrub or small tree up to 12 m high, all young parts with 
dense rufous tomentum. Leaves of the vegetative stems spirally arranged, impari- 
pinnate, multijugate, up to 60 (or more) X 32 cm; leaflets oblong, broadly 
elliptic or rotund, up to 16 X 7 cm, coriaceous, margin entire slightly revolute, 
base truncate to rounded, unequal, apex rounded, obtuse, or shortly and bluntly 
apiculate, mid-rib prominent, lateral veins usually clearly visible below, upper 
surface frequenty rugose. Leaves of the flowering branches smaller with fewer 
pinnae or more frequently unifoliolate, leaflets more rotund and with longer 
petioles. Inflorescence a cluster of specialized leafy branches forming a terminal 
crown; the main rays up to 60 cm long, bearing one or more rarely two opposite 
pairs of usually unifoliolate rotund leaves. 


Endemic to Borneo. Type: Sabah, Kinabalu, Clemens 33195 (BM). 


In forest and scrub from 600-2700 m, sometimes growing as an epiphyte in 
the crowns of trees. 


The bark is grey and smooth, the wood pale and soft, and the cut stems 
exude a yellowish or orange latex. This species is characterized by the rotund 
coriaceous leaves on the flowering branches. 


7. Arthrophyllum crassum Philipson, sp. nov. 


- Arbor parva in prima aetate tomento rufo. Folia imparipinnata, foliolis 
ellipticis oblongis vel lanceolatis. Inflorescentia ramis in proprium officium mutatis 
et in coronam termnialem dispositis, quater ramificantibus; bractea ovata 
plerumque acuta, rigidior alutaequa naturam habens, margine valde Tevoluta, 
nervis lateralibus obscurissimis. | 


306 Gardens’ Bulletin, Singapore — XXX (1977) 


Sparingly branched shrub or small tree up to 8 m high, all young parts with 
dense rufous tomentum. Leaves of the vegetative stems spirally arranged, impari- 
pinnate, multijugate, up to 100 (or more) X 50 cm, of the flowering branches 
smaller with fewer pinnae or more frequently unifoliolate; leaflets elliptic, oblong 
or lanceolate, up to 24 X 7.5 cm, coriaceous, margin entire, strongly revolute, 
base cuneate to rounded, often oblique, apex narrowed to an acute often caudate 
apiculum, mid-rib prominent, lateral veins faint to obscure. Inflorescence a cluster 
of specialized leafy branches forming a terminal crown; the main rays up to 70 cm 
long, bearing an opposite pair (or rarely more pairs) of unifoliolate or (less 
frequently) pinnate leaves. 


Endemic to Borneo. Type: Sarawak, Bako National Park, Chai $18020 
(Leiden). 


Swampy peat forest and heath woodland, both primary and disturbed, from 
sea level to about 150 m. Some fragmentary collections (Banying ak Nyudong 
S19404; Brunig S8720; Hewitt 770) from higher altitudes (1000 m) further 
inland may belong to this species, 


The leaves associated with the inflorescence are distinctively fleshy, have a 
strongly revolute margin and a smooth lower surface with indistinct lateral veins, 
and are ovate with rather acute apex. The leaflets of the pinnate leaves on the 
vegetative shoots are also rather leathery with indistinct lateral venation. The 
shrubs are occasionally epiphytic. 


8. Arthrophyllum pacificum Philipson, sp. nov. 


Arbor gracillis glabra, foliis imparipinnatis; foliolis plerumque ellipticis, c. 
8-10 x 4-5 cm, membranaceis Inflorescentia ramis in proprium officium mutatis 
et in coronam terminalem dispositis, quater ramificantibus. 


A slender tree to 14 m high, glabrous. Leaves of the vegetative stems spirally 
arranged, pinnate, multijugate, about 60 x 24 cm; of the flower-bearing branches 
smaller with fewer pinnae or simple; blade elliptic, oblong or avate, about 
8-10 x 4-5 cm, rather membranaceous, mafgin entire, very slightly revolute, base 
abruptly cuneate, often oblique, apex obtuse, acute, or slightly apiculate, Inflores- 
cence a whorl of specialized leafy branches forming a terminal crown to the 
vegetative shoots; the main rays 35 cm long bearing small pinnate leaves in 
Opposite pairs. 

Extending from the Moluccas to the Bismarck Archipelago. Type: Bismarck 
Archipelago, New Britain, Pomio subdistrict, Jsles et al NGF 34424 (Lae). 


Primary forest on slopes of mountains, attaining Nothofagus mossy forest, 
from 800-2000 m. Reported as common in most localities. 


All specimens from islands to the north of New Guinea are similar, in having 
more delicate foliage than O. macranthum from the mainland of New Guinea. 


9. Arthrophyllum diversifolium BI. Bijdr. Fl. Ned. Ind. (1826) 879. — A. 
javanicum Bl. l.c. — A. ellipticum Bl. 1.c. — Mormoraphis sumatrana Jack, ex 
Wall. Cat n. 4931 (1831). — Arthrophyllum blumeanum Zoll and Moritz, Syst. 
Verz. Pl. Zoll. (1845-6) 41. — A. ovalifolium Jungh. and de Vriese in Ned. 
Kruid. Arch. 1 (1846) 19. — A. ovatifolium Jungh. and de Vriese in Ann. Sci. 
Nat. 3, 6 (1846) 117. — A. dilatatum Miq. Fl. Ind. Bat. 1 (1856) 768. — A. 
diversifolium var. lanceolata Miq. Fl. Ind. Bat. Suppl. 1 (1860) 340. — A. 
blumeanum var. oblongatum Miq. in Ann, Mus. Bot. Lugd. -Bat. 1 (1863) 27; 
var. ellipticum Mig. |.c.; var. ovalifolium Miq. l.c. — A. borneense Baker in Kew 
Bull. (1896) 23. — A. congestum Ridley in Journ. Fed. Mal. St. Mus. 10 (1920) 
137. — A. havilandii Ridley in Kew Bull. (1933) 494. A. rufo-sepalum Ridley 
lic. (1946) 40. — A. rubiginosum Ridley l.c. 41. 


Arthrophyllum 307 


A small tree up to 14 m high, young parts with rufous tomentum. Leaves 
clustered at the end of the branches, spirally arranged on the vegetative shoots, 
imparipinnate or bipinnate (rarely tripinnate), multijugate, 150 cm long and 45 cm 
wide (wider in bipinnate leaves); usually in opposite pairs on the inflorescence- 
bearing branches and smaller with fewer pinnae or unifoliolate; leaflets ovate- 
oblong or elliptic, up to 24 x 11 cm + coriaceous or membranaceous, margin 
entire, slightly revolute, base truncate, rounded, or cuneate, often oblique, apex 
shortly acuminate. Inflorescence a cluster of specialized leafy branches forming a 
terminal crown which abscises after fruiting; the. main rays up to 150 cm long, 
bearing pinnate (or more rarely unifoliolate) leaves mostly in opposite pairs; the 
secondary rays up to c. 30 cm long, bearing mainly simple leaves in opposite pairs; 
tertiary rays (peduncles) about 5 cm long, articulate about the middle; pedicels 
+ 20, c. 5-12 mm long. 


From the Malay Peninsula and Sumatra to Java, Borneo and Celebes. Type: 
Java, Salak, Blume (Bogor). 


From sea-level to 1600 m, in a wide variety of habitats, from dry sandy soil 
to swampy humus. Occours in primary lowland and montane rain-forest and also 
in secondary forest, heath-forest and waste land. 


The very widespread A. diversifolium is variable in many characters, and many 
comprise a number of geographic subspecies, but no basis for this is apparent at 
present. Most individuals have the lower leaves simply imparipinnate, whereas 
others have bipinnate, or rarely tripinnate, leaves. The flowers and inflorescences 
of these forms appear to be identical, though raipd changes in the umbellules 
after flowering produce a deceptively distinctive appearance in specimens at 
different stages of development. Field experience over the whole range of the 
species will be required to understand this interesting leaf-polymorphism. In 
treating all forms as one species I am partly influenced by the fact that most 
authors who have been familiar with the plants in Java (where both forms occur) 
have regarded the complex as a single species (the fact that some authors have 
recognized the variant from Salak as a distinct species does not affect the problem 
of leaf polymorphism). 


Apart from the strikingly different leaf forms just discussed, certain local 
variants may eventually be shown to justify specific rank. A form growing on 
Salak (near Bogor) has often been regarded as distinct (see, for example, 
Hochreutiner in Candollea 2 (1925) 481, and Backer and Bakhuizen v.d .Brink, 
FI. Java 2 (1965) 169). Indeed this form is the basis of the name A. diversifolium. 
I retain this name in preference to the other two names published simultaneously 
by Blume because it has been most consistently adopted since it was first used in 
this comprehensive sense by Clarke. On the evidence available I do not consider 
the Salak plants any more distinctive than many other local variants. It might be 
considered that A. engganoense is also no more than another such variant, but its 
facies is so marked that specific rank appears justified. It is possible that Ridley 
was correct in distinguishing A. congestum, but the material is not good and 
appears inadequate to confirm specific status. Five collections from Brunei and a 
neighbouring district of Sarawak are all very alike and sufficiently distinct from 
both A. diversifolium and A. crassum to suggest that they represent a further 
species: these are Ashton S7840, Chew Wee-lek 981; Fuchs and Diederix 21182; 
and van Neil 3895 and 4312. For the present these five gatherings are tentatively 
retained as a form of A. diversifolium. Similarly, the two collections described by 
Ridley as A. rubiginosum and A. rufo-sepalum are based on collections which are 
not altogether typical of A. diversifolium, but which come closest to that species. 
In. the absence of more supporting material, it is advisable not to retain them as 
species. The first of these names (A. rubiginosum) has been widely used in 


308 Gardens’ Bulletin, Singapore — XXX (1977) 


identifications of Bornean ‘specimens, but the specimens concerned are either 
typical A. diversifolium or belong to the distinctive A. crassum. Specimens from 
Mt. Kinabalu described by Ridley as A. havilandii have bipinnate leaves, and 
appear to conform well with A. diversifolium. This form was again collected on 
Mt.- Kinabalu by Clemens and is also known from Sarawak (e.g. Morshidi 
S24074). | 


10. Arthophyllum macranthum Philipson, Bull. Brit. Mus. Nat. Hist. Bot. 1 
(1951) 18. 


Tree up to 25 m high, sparsely branched with leaves crowded at the ends of 
the branches, all young parts with dense rufous tomentum. Leaves of the vegeta- 
tive stems spirally arranged, imparipinnate, multijugate, up to 100 x 30 cm; of 
the flower-bearing branches smaller with fewer pinnae, or simple; petioles; leaflets 
ovate to oblong, up to 16 X 8 cm, coriaceous, margin entire, revolute, base 
rounded, truncate, or cordate, very rarely cuneate, often oblique, apex obtuse or 
bluntly apiculate. Inflorescence a whorl of specialized leafy branches forming a 
terminal crown; the main rays up to 60 cm long, bearing pinnate leaves often in 
opposite pairs; the secondary rays about 10-20 cm long, bearing simple leaves, 
usually in opposite pairs, and terminating in umbellules or compound umbellules; 
umbellules with about 10-15 flowers; pedicels stout 3-5 mm long to 10 mm or 
more in fruit. 


Endemic to New Guinea. Type: Morobe Distr. Boana, Clemens 8433 (A). 


Usually a sub-canopy tree of rain forest ranging from the lower montane 
zone to mossy subalpine woodland and scrubland. Occasional in second growth. 
Usually above 1000 m (up to 2700 m) but occurs on the coastal scarps of the 
Astrolabe range. 


Trees become very different in appearance when in flower or fruiting: the 
spiral pinnate foliage leaves are surmounted by tufts of branches which end in 
inflorescences and bear much smaller leaves. The foliage leaves are fleshy, leathery 
and glossy. The ripe fruit is purple and shining. The bark is grey, at first smooth 
with many leaf-scars and lenticels, but small longitudinal fissures develop. The 
cut branches exude a brown latex and a scent of celery. The soft wood is white 
or straw coloured. 


11. Arthrophyllum papyraceum Philipson, sp. nov. 


Frutex in prima aetate tomento rufo, mox glabrescens, Folia imparipinnata, 
foliolis circa 7; foliola tenuia, plana, c. 12-24 x 5-10 cm. Inflorescentia umbella 
terminalis composita radiis primariis paucis. 


A shrub rufous tomentose on the young parts. Leaves alternate, imparipinnate; 
leaflets about 7, membranaceous elliptic c. 12-24 x 5-10 cm, base broadly 
cuneate, apex finely acuminate, margin entire, slightly revolute. Inflorescence a 
terminal compound umbel; primary rays few (2), 2-3 cm long 2 mm wide, 
without bracts (caducous), secondary rays few (3) c. 13-18 mm long, articulated 
about the middle, ending in an umbellule of c. 10-12 flowers; pedicels 2-3 mm 
long slightly furfuraceous. 


Sumatra, Type: East Coast: Loendoet Concession, Koealoe, near Aek 
Sordang. Bartlett 6968 (Michigan). | 
- Collected in primary rain forest. 

_. - Known from a single collection (with no duplicates) this species resembles 


A. maingayi in its simple inflorescence, and the few pinnae of its foliage leaves. 
However, the large size of the leaflets precludes its inclusion in that species. 


Arthrophyllum 309 


12. Arthrophyllum maingayi Philipson, nom. nov. —- “A. pinnatum (Lam.) 
Clarke” in Hook, f. Fl. Brit. Ind. 2 (1879) 734, excluding synonyms. 


A low shrub or slender tree, rarely as high as 10 n, rufous tomentose on the 
very young parts, soon becoming glabrous. Leaves tufted at the end of the 
branches, spirally arranged, imparipinnate, usually with 5-7 leaflets (but up to 
15), up to 30 X 18 cm; leaflets elliptic or elliptic-lanceolate, + 8 xX 3.5 cm, 
rather thin, base cuneate or rounded, apex apiculate or caudate, margin entire, 
slightly revolute, lateral veins faint and obscure. Inflorescence consisting of a 
number of primary branches radiating from the end of a leafy shoot; the primary 
branches, often rather few, usually 3-6 cm long and devoid of leaves except for a 
few terminal simple or 3-foliolate leaves around the compound umbels, occasio- 
nally the branches bear pairs of opposite leaves when they may be up to 30 cm 
long; secondary rays c. 2-3 cm long; pedicels 4-10 cm long. 


Extending from Sumatra throughout the Malay Peninsula to Borneo. Type: 
Maingay 679 (SING). 

The name used by Clarke in Flora of British India has been in general use 
for this species. However, this name was based on Pannax pinnatum Lam. This 
in turn was based on the Rumphius name ‘“‘Scutellaria secunda’, which is quite a 
plant (Polyscias pinnata), Clarke therefore misapplied the name when using it for 
a Malay Peninsula Arthrophyllum, and a new name is necessary. 


The three species A. maingayi, A. montanum and A. alternifolium are similar 
in having simpler inflorescences than A. diversifolium. A. montanum can be 
distinguished from A. maingayi by its more leathery leaves with more prominent 
nervation, and by the more woody and leafy flower-bearing primary branches of 
the inflorescence. A. alternifolium is distinguished from both these species by its 
small coriaceous leaflets with inconspicuous nervation. Most specimens can be 
readily distinguished, but a few can be asigned to a species only doubtfully, 
usually because the material is inadequate. This is usually due to the junction 
between the vegetative (spiral phyllotaxtic) shoots and the flower-bearing branches 
being omitted. 


13. Arthrophyllum kjelbergii Philipson, sp. nov. 


Arbor parva in prima aetate tomento rufo. Folia imparipinnata, foliolis 
ellipticis vel ovate-oblongis, ad 8 X 3.3 cm, chartaceis. Inflorescentia ramis in 
proprium officium mutatis et in coronam terminalem dispositis, ter ramificantibus. 


A small tree, 10 m high, branches c. 1 cm diam., the young parts covered in 
rufous tomentum, Leaves clustered towards the ends of the branches, spirally 
arranged on the vegetative shoots, imparipinnate, multijugate, 25-30 x 16 cm; 
leaflets elliptic or ovate-oblong, up to 8 X 3.3 cm, chartaceous, margin entire 
minutely revolute, base rounded to cuneate often oblique, apex tapered to a blunt 
apiculum, or rounded and mucronate. Inflorescence a terminal cluster of c. 5-10 
specialized branches; primary rays c. 15-20 cm long, 1.5-2.0 mm wide bearing 
near the middle an opposite pair of simple leaves or sometimes trifoliate leaves, 
and 2-3 similar leaves below the terminal cluster of secondary rays; secondary 
rays c. 12, c. 2.5—3.5 cm long, terminating in an umbellule of about 12 flowers. 


Endemic to Celebes. Type: South-east Celebes: near Kendari, Sangona, 
Kjellberg 1145 (Bogor). 


In primary rain forest 50-150 m. 
The small leaves and relatively simple inflorescences are distinctive. 
14. Arthrophyllum cenabrei Merr. in Philip. Journ. Sci. 20 (1922) 417. 


A glabrous tree c. 10 m high, the ultimate branches c. 5 mm diam. Upper 
leaves pinnate, up to 10 cm long, leaflets mostly 5, sometimes 3, or the uppermost 


Gardens’ 


310 


Bulletin, Singapore — XXX (1977) ' 


‘UINSSDID *p 
‘8 ‘winjuayjoo -y “Lt “(£ x) sprig 
‘WMNnjudpsonUu 
V ‘Ql Swnyofissaaip "py ‘cy 
‘asuaouvs3ua “p ‘py “uinupiusayo 
Vv ‘Il (= x= ae) somypoquya 


ZY 


‘81 


‘ee 


“LI ‘ss1y 


—-€[ ‘sary 


V ‘Il 
eB 3 


‘wWaaqyaly “yp 
‘ipsum “p 


‘ 


Ol 


‘ 


L 


(¢/T x |e) soavoy o8er0.7 ‘TI-L  “Sd1q 


‘1uOolysD “py ‘Z] ‘uinyofiusajjo 
‘asuadv3jnd “p ‘6 ‘wnupjuow 


Arthrophyllum 311 


reduced to simple leaflets, the rhachis and petiole c, 4 cm long, the leaflets mostly 
elliptic, 4.5-6 xX 2.5-3.5 cm, chartaceous to subcoriaceous, very shortly and 
obtusely acuminate, base acute, brownish olivaceous and slightly shining when 
dry. Peduncels c. 4 cm long, umbellately arranged at the tops of the branchlets, 
usually however with solidary inflorescences in the axils of the uppermost leaves, 
thus forming a somewhat leafy inflorescence. 


Endemic to the Philippines. Type: Cebu, Cenabre and de la Cruz FB28343 
(n.v.). 


On slopes at 600 m. 


No specimen of this species has been located. The above description is taken 
from Merrill’s original account. In placing this species in the key, it has been 
assumed that the inflorescence branching is relatively simple. 


15. Arthrophyllum pulgarense Elmer in Leaflets Philipp. Bot. 7 (1915) 2551. 


A small tree, branches c. 1 cm diam., young parts with red tomentum, 
becoming glabrous except on the ovaries. Leaves clustered towards the ends of 
the branches, spirally arranged on the vegetative branches, imparipinnate, about 
6 pairs of leaflets, c. 22 X 8 cm; leaflets elliptic to rotund, c. 4 xX 2.8, coriaceous, 
margin entire, revolute, rounded to broadly cuneate, apex rounded or abruptly 
tapered to a short obtuse apiculum, Inflorescence a terminal cluster of specialized 
branches; primary rays c. 6-10 cm long, 3-4 mm wide, bearing near the middle 
an opposite pair of simple rotund leaves, and with a whorl of similar leaves below 
the terminal cluster of secondary rays; secondary rays c. 6-8, 2-4 cm long 
terminating in an umbellule of about 8-12 flowers, bracts caducous. 


Endemic to the Philippines. Type: Palawan, Elmer 13193 (PNH). 
Common in montane forests on Mt Pulgar. 
The coriaceous small often rotund leaflets are characteristic. 


16. Arthrophyllum montanum Ridley in Journ. Fed. Mal. St. Mus. 4 (1909) 
24. — A. nitidum Ridley l.c. 7 (1916) 42. — R. ovatum Ridley l.c. 42. 


Shrub or small tree to 6 m, unbranched or sparingly branched, rufous tomen- 
tose on the young parts, becoming glabrous. Leaves tufted at the ends of the 
branches, spirally arranged, imapripinnate, multijugate, about 30-55 x 12-22 cm; 
leaflets elliptic or oblong 6-10 x 2.5-4 cm, coriaceous or charactaceous, base 
cuneate, apex with a short blunt apiculum, margin entire, revolute, the few 
principal lateral veins usually rather prominent; the leaves associated with the 
umbels usually unifoliolate, broadly elliptic to rotund. Inflorescences on specialized 
leafy branches either in terminal clusters or axillary in the upper leaves; the 
branches 10-30 cm long, leaves mostly simple in opposite pairs, the branches 
ending in a whorl of simple leaves surrounding a compound umbel. 


Malaya Peninsula. Type: Pahang, Ridley 13687 (SING). 
Primary forest ascending to montane zone, and in second growth, 250-1500 m. 


Similar to A. pinnatum but distinguished by the more leathery leaves and by 
the leafy inflorescence branches which are usually absent in A. pinnatum. 


17. Arthrophyllum alternifolium Ridley Fl. Mal. Pen (1922) 886. — A. 
pinnatum Clarke in Hook. f. Fl. Brit. Ind. 2 (1897) 734, pro parte; King in 


Journ. As. Soc. Bengal 67 (1898) 59, pro parte. — A. alternifolium Clarke l.c., 
in synon, 


A slender sparingly branched shrub, to 2 m high, rufous tomentose on the 
young parts, becoming | glabrous, Leaves tufted at the ends of the branches, 
spirally arranged, imparipinnate, multijugate, about 20-25 (30) x 9-12 (15) cm; 


312 Gardens’ Bulletin, Singapore — XXX (1977) 


leaflets ovate, elliptic or lanceolate, 3.5-4 (6) xX 0.9-2 (2.3) cm, coriaceous, base 
cuneate, apex acuminate to caudate, obtuse, margin entire, revolute, veins obscure; 
the leaves associated with the umbels (if any) reduced, with fewer leaflets or 
unifoliate, sometimes broadly ovate. Inflorescence usually a terminal compound 
umbel, occasionally a whorl of leafy branches (5-14 cm long) each ending in a 
compound umbel. 

Malay Peninsula. Type: Malacca, Griffith 2676 (SING). 

In shady montane forest, with Rhododendron and Dacrydium, 3000 ft and 
above. 


The small coriaceous, often apiculate, leaflets are characteristic, Although 
collected most frequently on Mt Ophir it occurs on other high ridges in the 
southern Peninsula. 


INSUFFICIENTLY KNOWN SPECIES 
A, reticulatum Bl. ex Miq. in Ann. Mus. Bot. Lugd. -Bot. 1 (1863) 27. 


The original description was based on foliage leaves only. Their place of 
origin was uncertain. Since I have seen no specimen identified as this species, its 
identity cannot be decided. 


A. serratifollum Miq. Fl. Ind. Bat. Suppl. (1860) 341. 


The original description was based on foliage leaves only. I have seen no 
specimen identified as this species, but since all other members of the genus have 
entire margins to the leaflets, [ doubt if this species would prove to be a true 
Arthrophyllum. 


[ wish to thank Dr E. Edgar of Botany Division, D.S.I.R., Christchurch for 
writing the Latin diagnoses. 


PUBLICATIONS OF THE BOTANIC GARDENS 
SINGAPORE 


1 The Agricultural Bulletin of the Malay Peninsula (Series I). 
Only Nos. 3, 5, 7, 8 and 9 available, at 20 cents each. 


2. The Agricultural Bulletin of the Straits and F.M.S. (Series ID). 


Vols. 1-10, 1901-1911, monthly issues. 
Many parts available. 
Price: $5 per volume, 50 cents per part. 


3. The Gardens’ Bulletin, Straits Settlements (Series LID). 


Vols. 1-11, 1912-1947. 
Vol. | (1-5) January-May 1912 issued under title of Agricultural Bulletin 
of the Straits & F.M.S. 


Prices on application. 


4 The Gardens’ Bulletin, Singapore (Series TV). 
Vols. 12-27, 1949-1971. Vol. 28 Pt. I in print. 


Price: Vol. 13 Pt. I (New impression) $12 per copy, $20 per vol. 

Vols, 12 & 14 $13 per vol. Vol. 15 $20 per vol. Vols. 16-25 $25 
vol. Individual parts vary in price. 
Wonrco (it. t Sid; at. ea Slaps 27 APt. 1 S18; Pt. I $8.50); 
28 (Pt. I $18); 29 ($30). In press: The Freshwater Swamp-forest 
of S. Johore and Singapore by E. J. H. Corner (Gdns’ Bull. 
Suppl. 1). 


5. Materials for a Flora of the Malay Peninsula, Monocotyledons. 
Parts 1, 2 and 3 remain available. 
Price: $10 per set, $5 per part. 

6. Annual Reports. 
1909-1972. 


7. (a) Malayan Orchid Hybrids by M. R. Henderson and G. H. Addison. $15. 
(b) Malayan Orchid Hybrids, Supplement I by G. H. Addison. $21. 


8. A Revised Flora of Malaya. 


(a) Vol. 1, Orchids, by R. E. Holttum. $30 (3rd ed. 1972 Impr.). 
(b) Vol. 2, Ferns, by R. E. Holttum. $20 (2nd ed., 1968). 
(c) Vol. 3, Grasses, by tH. B. Gilliland. $30 (1971). 


9. Boletus in Malaysia by E. J. H. Corner. $50 (1972). 


Items 1-6 obtainable from the Commissioner, Parks & Recreation Department, 
Botanic Gardens, Cluny Road, Singapore 10. 

Items 7-9 obtainable from Singapore Nationa! Printers (Pte) Ltd, Upper Serangoon 
Road, Singapore 13. Ri 


Prices quoted are in Singapore Dollars 
Overseas postage Is extra 


if 


Pa ee 
Satan 


+e 


ee ys 
as eae 
eee, IPE nnn, 
pe tet ep eae te rey 
ah ~~ SL ye 

SRN re WS ‘ Saal E ; ~ ' 

ee SS ON ne Sy pegstit DA ast aie 2 

SS a SS en Ln ~ Owe “Te Sr. Se etys 

Pag en agen) 7 pi din 2 

= I nn ~ : 
. 2 ary 

a Ney. > 


biped dh th ok ee 
~N 6e 


a > eT tate i le 
“<~ eee ee, — 


ap aaon ate Artes” 
hie ne en eee ewe, Pe . 
© » es we “ee a 


ee ee ey 


OE Rs 20°4, : 
tind nile hsdpa tn Seer Ree